On the Actinopterygian Fish Fauna
(Upper Cretaceous: Campanian)
from the Kristianstad Basin,
Southern Sweden
Mohamad Bazzi
Actinopterygian remains have been recovered from Upper
Cretaceous (uppermost lower to lowermost upper Campanian)
marine strata of the Kristianstad Basin, southern Sweden. This
is the first record of Upper Cretaceous bony fish from the
Fennoscandian shield. The fauna consists of higher taxa
including Pachycormiformes (Pachycormidae), Elopiformes
(Pachyrhizodontidae), Pycnodontiformes (Pycnodontidae),
Aulopiformes
(Enchodontidae),
Ichthyodectiformes
(Ichthyodectidae), and indeterminable teleost fish. The
ichthyofauna comprises a relative vast number of
disarticulated meso-to macroscopic bony elements (e.g.
vertebrae, isolated teeth, cycloid scales, coprolites, otolithes
and fin spines). All major groups are so far only recognized by
a few numbers of isolated dental remains suggesting major
taphonomic losses prior to preservation. The total amount of
recovered bony fish elements indicates the presence of a
diverse teleostean/actinopterygian fauna, although taxonomic
assessments of the material are severely hampered by the low
preservational degree. In addition, previous works on the
vertebrate assemblage of the Kristianstad Basin have resulted
in major collection biases towards larger marine vertebrates
which has led to an under-representation of the
actinopterygian fauna. The ichthyofauna of the Kristianstad
Basin bears compositional resemblance towards those in the
North American Western Interior Seaway suggesting
palaeobiogeographical communication.
Uppsala universitet, Institutionen för geovetenskaper
Kandidatexamen i Geovetenskap, 180 hp
Självständigt arbete i geovetenskap, 15 hp
Tryckt hos Institutionen för geovetenskaper
Geotryckeriet, Uppsala universitet, Uppsala, 2014.
Självständigt arbete Nr 88
On the Actinopterygian Fish Fauna
(Upper Cretaceous: Campanian)
from the Kristianstad Basin,
Southern Sweden
Mohamad Bazzi
Självständigt arbete Nr 88
On the Actinopterygian Fish Fauna
(Upper Cretaceous: Campanian)
from the Kristianstad Basin,
Southern Sweden
Mohamad Bazzi
Supervisors: Benjamin P. Kear and Elisabeth Einarsson
Sammanfattning
Material tillhörande fossila strålfeniga benfiskar har insamlats från marina avlagringar
av övre krittid (senast tidigcampan till tidig sencampan) från Kristianstadbassängen,
södra Sverige. Detta material representerar det första fossila registret av benfiskar
från övre kritavlagringar inom den Fennoskandiska skölden. Fiskfaunan utgörs av
högre taxonomiska grupper såsom Pachycormiformer (Pachycormidae),
tarponartade fiskar (Pachyrhizodontidae), Pycnodontiformer (Pycnodontidae),
laxtobisartade fiskar (Enchodontidae), Ichthyodectiformer (Ichthyodectidae) samt
oidentifierbara teloster. Det fossila fisk materialet utgörs av en mängd disartikulerade
meso- till makroskopiska element (ex. kotor, tänder, cykloidfjäll, koproliter, otoliter
och fentaggar). Alla identifierade fiskgrupper som hittills dokumenterats är
representerade enbart av isolerade tänder, något som indikerar på tafonomisk förlust
tillföljd av bevaring. Det totala benfisk material som insamlats indikerar på att faunan
var mångfaldig dock hindras taxonomiska bedömningar från att göras på grund av
det låga bevaringstillstånd materialet uppvisar. Tidigare arbeten som utförts på
fossila ryggradsdjur från Kristianstadbassängen har resulterat i partisk insamling i
huvudsak gentemot större marina vertebrater som i sin tur lett till en tydlig
underrepresentation av den strålfeniga fiskfaunan. Fiskfaunan i
Kristianstadbassängen är väldigt lik den som återfinns i den Nordamerikanska
”Western Interior Seaway” något som indikerar paleobiogeografisk kommunikation.
Abstract
Actinopterygian remains have been recovered from Upper Cretaceous (uppermost
lower to lowermost upper Campanian) marine strata of the Kristianstad Basin,
southern Sweden. This is the first record of Upper Cretaceous bony fish from the
Fennoscandian shield. The fauna consists of higher taxa including Pachycormiformes
(Pachycormidae), Elopiformes (Pachyrhizodontidae), Pycnodontiformes
(Pycnodontidae), Aulopiformes (Enchodontidae), Ichthyodectiformes
(Ichthyodectidae), and indeterminable teleost fish. The ichthyofauna comprises a
relative vast number of disarticulated meso-to macroscopic bony elements (e.g.
vertebrae, isolated teeth, cycloid scales, coprolites, otolithes and fin spines). All major
groups are so far only recognized by a few numbers of isolated dental remains
suggesting major taphonomic losses prior to preservation. The total amount of
recovered bony fish elements indicates the presence of a diverse
teleostean/actinopterygian fauna, although taxonomic assessments of the material
are severely hampered by the low preservational degree. In addition, previous works
on the vertebrate assemblage of the Kristianstad Basin have resulted in major
collection biases towards larger marine vertebrates which has led to an underrepresentation of the actinopterygian fauna. The ichthyofauna of the Kristianstad
Basin bears compositional resemblance towards those in the North American
Western Interior Seaway suggesting palaeobiogeographical communication.
Key words: Upper Cretaceous; Campanian; Osteichthyans; Actinopterygian;
Teleostei; Palaeoecology; Taphonomy; Kristianstad Basin; Åsen; Southern Sweden
i
ii
Table of content
1. Introduction ........................................................................................................... 1
2. Previous research................................................................................................. 2
3. Stratigraphy and geological setting .................................................................... 2
4. Localities and associated vertebrate fauna ....................................................... 4
4.1 Åsen .................................................................................................................. 4
4.2 Ivö Klack ............................................................................................................ 4
4.3 Balsvik ............................................................................................................... 5
4.4 Ignaberga (new quarry) ..................................................................................... 5
4.5 Ullstorp .............................................................................................................. 6
5. Material & method................................................................................................. 6
6. Institutional abbreviations ................................................................................... 7
7. Systematic palaeontology ................................................................................... 7
8. Discussion .......................................................................................................... 16
8.1 Palaeoecology of the actinopterygian assemblage ......................................... 16
8.2 Pycnodontiformes ............................................................................................ 17
8.3 Aulopiformes ................................................................................................... 18
8.4 Ichthyodectiformes .......................................................................................... 18
8.5 Pachycormiformes ........................................................................................... 19
8.6 Elopiformes ..................................................................................................... 19
9. Taphonomic and palaeoenvironmental implications ...................................... 19
10. Conclusions ...................................................................................................... 23
11. Acknowledgments ............................................................................................ 24
12. References ........................................................................................................ 24
13. Appendices ....................................................................................................... 33
iii
iv
1. Introduction
Upper Cretaceous (Campanian) strata from the Kristianstad Basin, northeastern
Scania, Sweden, has yielded a plethora of fossil organisms. The variety of fossil
vertebrate components from the different fossil localities may be referred to as a
microvertebrate assemblage. These comprise a diverse array of species including a
quantity of chondrichthyan (members, from both the Elasmobranch and Holocephali
clade) elements (e.g. mostly teeth). Also, osteichthyans (bony fish) and marine
tetrapods, such as plesiosaurs, mosasaurs, aquatic birds, dyrosaurid crocodilians,
and unidentified taxa of cheloniid sea turtles were major faunal components of this
ancient marine ecosystem (Davis, 1890; Persson, 1959, 1963; Siverson, 1992a;
Lindgren, 2004, 2005; Rees & Lindgren, 2005; Sørensen et al., 2013). In addition a
high density and diversity of macroinvertebrates such as echinoids, belemnites,
brachiopods, bivalves (predominantly oysters) and bryozoans (Surlyk & Christensen,
1974; Christensen, 1975; Sørensen & Surlyk, 2008, 2010, 2011; Surlyk & Sørensen,
2010; Sørensen et al., 2011, 2012) have been identified. Furthermore, the
Kristianstad Basin has yielded continental elements such as dinosaurs represented
by isolated dental and caudal remains of neoceratopsian ornithischians (Lindgren et
al., 2007).
Here, I report upon the occurrence of marine actinopterygian and teleost
fossil fish from Upper Cretaceous (Campanian) strata from the Kristianstad Basin. In
comparison to the extensively documented records of other vertebrate groups (e.g.
Sørensen et al., 2013), almost nothing has been said about the bony fish
assemblage. Thus, the recovery of fossil fish, with a Campanian affinity, consisting
mainly of marine actinopterygians including pycnodontiforms (Anomoeodus
subclavatus Agassiz, 1833), pachycormids (Protosphyraena sp. Leidy, 1857a),
enchodontids (Enchodus gladidus Cope, 1872b), elopiformes (cf. Pachyrhizodus sp.
Dixon, 1850), ichthyodectids (indet. Ichthyodectidae Crook, 1892), and
indeterminable teleost fish is of significance, because it provides new account of
Upper Cretaceous osteichthyan ecology from the region. In addition, this paper aims
to provide new information about the taxonomic occurrence of fossil fish from
southern Sweden. It represents the first study on the subject and complements
previous palaeoecological studies (e.g. Sørensen et al., 2012, 2013) of the
Kristianstad Basin.
On the basis of previous work (e.g. Siverson, 1992a, b, 1993, 1995;
Rees, 1999) Sørensen et al., (2013) managed to report a total of thirty-eight
chondrichthyan species, from uppermost lower Campanian strata in the Kristianstad
Basin. Verily this attests to a highly productive ecosystem containing an array of
selachian taxa (Siverson, 1992a) and larger squamate species (e.g. Lindgren &
Siverson, 2002).
There are also reasons to suggest that actinopterygian fish occupied a
broad range within the basin. The nature of preservation of actinopterygian remains
is however very poor and problematic in terms of taxonomy, but still they form a vital
as well as a prominent portion of this Upper Cretaceous inner shelf- shallow marine
environment.
1
2. Previous research
Davis (1890) described and reported on several bony fish taxa from the Mesozoic
and Early Cenozoic (Danian) of Scania, southern Sweden. Among the reported
specimens, were pycnodontiformes (“Coelodus” subclavatus, p. 415-416, pl. 42, fig.
16-18), beryciformes (Hoplopteryx lundensis Davis, 1890, p. 417-420, pl. 43, fig. 1-3;
Hoplopteryx sp., p. 420-421, pl. 42, fig. 19-20; Hoplopteryx minor Davis, 1890, p.
421-422, pl. 45, fig. 3-4; Berycopsis lindstromi Davis, 1890, p. 422-423, pl. 44, fig. 11a), aulopiformes (Enchodus sp., p. 424, not figured; Dercetis limhamnensis Davis,
1890, p. 431-432, pl. 45, fig. 1-2 ), lampridiformes (Bathysoma lutkeni Davis, 1890, p.
424-427, pl. 46, fig. 1-7), and clupeiformes (Clupea lundgreni Davis, 1890, p. 427428, pl. 45, fig. 5). Most of these specimens were reported from the Limnhamn quarry
and supposedly also from Köpinge, with the exception of the pycnodontiformes being
reported from the Ignaberga locality. Siverson (1992a) raised doubts and a proper
revision concerning the many locality errors found in Davis (1890) paper on the
Scandinavian fossil fish. It was also concluded in the same paper that most selachian
lamniform sharks figured by Davis (1890) were erroneously taxonomically designated
(Siverson, 1992a). No efforts have previously been made to confirm the true
geographic occurrence of the osteichthyan assemblage reported by Davis (1890).
Unfortunately, a proper reevaluation of the locality data and bony fish taxonomy
stated by Davis (1890) is not within the scope of the present paper. Although, it is
confirmed here that pycnodontiforms have been encountered at Ignaberga and those
dental remains figured by Davis (1890, p. 415-416, pl. 42, fig. 16-18) are correctly
identified as pycnodontiforms.
3. Stratigraphy and geological setting
Erlström & Gabrielson (1986, 1992), Lidmar-Bergström (1982), Bergström &
Sundquist (1978) and Christensen (1975, 1998) have provided detailed accounts of
geology and stratigraphy (including maps) for the Kristianstad Basin. In brief
summation, the Kristianstad Basin consists of a deeply weathered, crystalline
basement (Precambrian in age) that manifests a very uneven topography (Bergström
& Sundquist, 1978; Lidmar-Bergström, 1982). The basin is located in the province of
northeastern Scania, southern Sweden and is among six areas in Sweden were
Upper Cretaceous deposits are known (Bergström & Sundquist, 1978; Siverson,
1992a). During the Upper Cretaceous, the Kristianstad Basin was situated at a
palaeolatitude of 50° N (Smith et al., 1994), which puts the basin at the NE margin of
the former epeiric Chalk Sea, which during this interval covered a great portion of
northwestern Europe (Surlyk and Sørensen, 2010; Sørensen et al., 2013). The
climatic condition during the uppermost lower Campanian is considered to have been
between warm temperate to subtropical (Siverson, 1992a; Surlyk, 1997; Surlyk &
Sørensen, 2010). The global sea level in the Upper Cretaceous was 75-100 m higher
than it is today (Kominz et al., 2008), and as a result of repeated transgressive
phases, the area was transformed into an archipelagic environment surrounded by
low-lying islands and peninsulas (Surlyk & Christensen, 1974). As current
topography indicates, the crystalline basement forms two half grabens which are
intercepted to the south by the Nävlingåsen and Linderödåsen horst ridges (Lindgren
et al., 2007; Einarsson et al., 2010). The Upper Cretaceous marine succession of the
Kristianstad Basin is < 200 m thick and consists predominantly of detrial carbonates,
sand and silt (Surlyk & Sørensen, 2010).
2
Figure 1. Stratigraphy, age and associated fish fauna from the different horizons at the Åsen
locality. Black dots (Indet. Pycnodontidae), grey dot (Anomoeodus subclavatus), triangle with
a black dot in the center (Indet. Ichthyodectidae), triangle with three black dots (cf. Enchodus
sp.), rhombus (cf. Pachyrhizodus sp.), and M (Indet. Teleost A.).
Figure 2. Schematic geological map of the fossil localities of the Kristianstad Basin,
Skåne/Scania, southern Sweden. The Ullstorp site is not shown as an enlarged section in the
map; however it has been given its relative geographical position in the enlargement of
Scania and lies within the vicinity of Ignaberga (B). The scale bar shown in A-D is 1 km (after
Bergström & Sundquist, 1978; Bergström, 1983a, b; Kornfält & Bergström, 1989).
3
4. Localities and associated vertebrate fauna
4.1 Åsen
The Åsen locality (N56°08´56.1´´, E14°29´56.0´´; WGS84; map sheet: Karlshamn 3E
NV, co-ordinates: UTM VC 690 229; Fig. 2A) is an abandoned clay pit, currently used
as a rubbish dump (Sørensen et al., 2013). The locality is situated in the northeastern
corner of the Kristianstad Basin and has produced the best paleontological record of
marine macrovertebrates from Upper Cretaceous deposits in Sweden. Åsen has also
produced exceptionally persevered plant material, and is regarded as a Upper
Cretaceous Lagerstätte (Friis & Skarby, 1981; Siverson, 1992a).The Åsen
stratigraphic succession is marine in origin and is divided into biostratigraphic zones,
informally named the Belemnellocamax mammillatus (uppermost lower Campanian)
and the Belemnellocamax balsvikensis (lowermost upper Campanian) belemnite
biozones (Christensen, 1975). The stratigraphic marine succession is made up of an
approximately 3.5 m thick greensand unit which is well stratified, glaciotectonized and
unconsolidated and overlies fluvio-lacustrine clayey and argillaceous deposits with
upper Santonian and/or lower Campanian affiliations (Friis & Skarby, 1981; Siverson,
1992a).
The vertebrate fauna reported from Åsen comprises thirty-six shark, six
bony fish, five ray, six mosasaur and one dinosaur species (Sørensen et al., 2013:
table 2, and this paper). Also, polycotylid and elasmosaurid plesiosaurs (Persson,
1959; Einarsson et al., 2010), aquatic birds, (J. Lindgren, unpublished data)
chimaeroids (Ischyodus sp. personal observation) and turtle remains have been
recovered. The bony fish materials from Åsen makeup the greatest portion of the
total teleost record from the Kristianstad Basin but are represented only by
disarticulated elements (e.g. vertebral centra and isolated worn teeth). Based on
personal observations, fine spines and mechanically disintegrated skeletal bony fish
elements constitute the bulk of the vertebrate fossil sample from Åsen. Altogether,
bony fish components are presumably equally abundant as cartilaginous fish
elements; however taxonomic assessments are of such material highly problematic
and therefore only diagnostic dental remains have been considered.
4.2 Ivö Klack
Ivö Klack (N56°08´21.6´´, E14°24´05.8´´; map sheet: Karlshamn 3E SV, coordinates:
UTM VC 631 222; Fig. 2D) also referred to as Blaksudden in older literature is an
abandoned kaolin and limestone quarry situated along the northern slope of the
Island of Ivö (Surlyk & Christensen, 1974; Siverson, 1992a). The crystalline
basement of the Ivö Klack locality formed an island in an archipelagic environment at
the time of uppermost lower Campanian. The surrounding sea contained a high
density and diversity of marine organisms (Siverson, 1992a; Surlyk & Sørensen,
2010; Sørensen et al., 2013). Due to the transgressive pulses of the Upper
Cretaceous Sea, a steeply, irregular shaped rocky shore emerged at Ivö island as a
result of wave action sweeping away the deeply weathered gneiss basement, leaving
a boulder strewn and hummocky rocky coast (Surlyk & Sørensen, 2010; Sørensen &
Surlyk, 2011). The shore formed part of the intertidal and subtidal zones with an
estimated water depth reaching nearly 5 m (Sørensen et al., 2012). The Cretaceous
stratigraphic succession at the Ivö Klack locality is made up by a 3-4 m thick
nonfossiliferous quarts sand unit that is overlain by a 22-24 m thick coarse-grained
4
marine carbonate unit which is composed of bivalvian skeletal components,
echinoids, bryozoans and brachiopods (Siverson, 1992a). The fossil components
found in the coarse-grained marine carbonate strata is predominantly made up by
oysters that formed banks along the shoreline of the island at the time of the Upper
Cretaceous and belongs to the uppermost lower Campanian Belemnellocamax
mammillatus belemnite biozone, c. 81-80 myr (Sørensen et al., 2010, 2012).
The Ivö Klack bony fish specimens described herein are part of an old
collection that has been loaned from the Department of Palaeozoology (RM PZ),
Stockholm, Sweden (Appendix 1.2). The marine vertebrate fauna known so far from
Ivö Klack comprises in total two bony fish, one ray, twenty three shark, six
mosasaur, four plesiosaur, three aquatic bird and one crocodilian species (Sørensen
et al., 2013: table 2, and this paper). The invertebrate fauna of Ivö has recently been
extensively documented and comprises > 60 bivalvian, eight zooxanthellate and
three azooxanthellate coral, seventeen gastropod, eighteen echinoid and sixteen
asteroid, four belemnite and one ammonite, and twenty-five tube-dwelling polychaete
species (Sørensen & Surlyk, 2010, 2011; Surlyk & Sørensen, 2010; Sørensen et al.,
2011, 2012).
4.3 Balsvik
The currently abandoned Balsvik quarry (N56°05´149´´, E14°14´115´´; map sheet:
Karlshamn 3E SV, coordinates: UTM VC 519157, Fig. 2C) exposes a succession
made up by conglomerates at its base in which the informal lowermost upper
Campanian Belemnellocamax balsvikensis and Belemnitella mucronata biozones are
present (Christensen, 1975, 1998). It is followed by an approximately 6 m thick unit
comprising fine-grained calcarenite yielding Belemnitella mucronata (Christensen,
1998). The upper Campanian sequence is terminated by a discontinuity surface
indicating a break in sedimentation (i.e. hiatus) and is immediately succeeded by a 3
m thick unit of calcisiltite and fine-grained calcarenite of Maastrichtian age
(Christensen, 1998; Siverson & Cappeta, 2001). The onset of the Maastrichtian in
northern Europe is marked by the appearance of Belemnella lanceolata (Christensen
et al., 2000).
The vertebrate fauna reported from the Balsvik quarry comprises thirty
selachian (Siverson, 1993b), and one mosasaur species from B. lanceolata
(Lindgren, 2004). Also two bony fish taxa have been recovered from unknown
horizon.
4.4 Ignaberga (new quarry)
The Ignaberga locality (N56°06´50´´, E13°51´24´´; map sheet: Kristianstad 3D SO,
coordinates: UTM VC 288 195, Fig. 2B) currently comprises two limestone quarries.
Carbonate deposits of uppermost lower Campanian in age are exposed at the new
quarry (Christensen, 1975). Well stratified skeletal sands containing breakdown
products from the adjacent gneiss horst (i.e. Nävlingeåsen) are usually observed in
exposed sediments at Ignaberga (Christensen, 1975; Birkelund & Bromley, 1979).
These Upper Cretaceous deposits are highly fossiliferous and comprise a variety of
fossil fragments belonging to bivalves, echinoids and bryozoans (Sørensen et al.,
2013).
The marine vertebrate assemblage reported from Ignaberga comprises
so far only one osteichthyan, twenty eight shark, two plesiosaur and three mosasaur
5
species (Sørensen et al., 2013: table 2 and this paper). The bony fish sample from
Ignaberga comprises only one specimen that belongs to Anomoeodus subclavatus.
4.5 Ullstorp
Ullstorp (N 56°2´44.4´´, E13° 57´20.9´´; map sheet: Kristianstad 3D SO, coordinates:
UTM VC 352 137) is situated 9, 9 km SE of Ignaberga and comprises five smaller
abandoned quarries which are closely positioned to the Nävlingeåsen to the south
(Bergström & Sundquist, 1978; Erlström & Gabrielson, 1985). These quarries have
been numbered from 1-5 (see Christensen, 1975; Erlström & Gabrielson, 1985).
Unfortunately bony fish specimens described herein cannot be ascribed to a specific
quarry. For a more detailed account of the geology of Ullstorp see Christensen
(1975) and Erlström & Gabrielson (1985).
Previously documented marine vertebrates from Ullstorp comprise nine
chondrichthyan and one crocodylian species (Siverson, 1992a). In addition, two
actinopterygian taxa and one unidentified pycnodontiform species have been
recovered. Also, incisiform and fang-like teeth of uncertain systematic affinity are
recorded at Ullstorp (Appendix 4.3).
5. Material & method
The Cretaceous fossil fish remains recovered to date from Scania consist of several
hundreds if not thousands of specimens. Most of these have been collected directly
from the Åsen locality (Appendix 1.1). The state of preservation ranges between
highly fragmented and damaged to well preserved, yet partly articulated masses and
bits. Also, dental remains attributable to pycnodontiforms (PMU 27255, PMU 27256,
PMU 27257; Fig. 3A-E) have been revised and included in this paper and are
currently housed at the Museum of Evolution (PMU), Uppsala, Sweden. Additional
fossil fish remains from Ullstorp and Åsen were donated from a private collector and
have been included in the collection at the Museum of Evolution in Uppsala.
The dominant portion of the fossil fish material described herein was
collected during field excavations at the Åsen locality in 2011. The research
excavations at the Åsen site, forms a part of an ongoing research project, which has
facilitated a reassessment of the marine paleoecology of the Kristianstad Basin (E,
Einarsson et al., unpublished data). The materials were extracted from the poorly
consolidated marine sediments by means of direct field sieving (using a mesh size of
~ 2.5 mm). In total all sampled horizons at the Åsen locality (sieved sediments 6655.
5 Kg), managed to produce an abundant fossil material. The laboratory components
of this project consisted of sorting the materials, examining the material under a
binocular microscope and establish measurements of each individual specimen by
using a ruler. Out of 557 examined bony fish components (e.g. fin spines, vertebral
centra and teeth) from Åsen only those with diagnostic features were considered.
This concludes that the dominant portion of the fossil specimens from the Åsen
locality lacked diagnostic features, hence left those without an osteological and/or
taxonomical identification. In turn to avoid problems of parataxonomy, only
specimens with sufficient diagnostic traits have been subject to taxonomical
designation, hence left those simply generically indeterminable aside, with the
exception of frequently encountered dental remains that are of ecomorphological
value. The palaeontological record of extinct actinopterygian taxa from the
Kristianstad Basin are represented by teeth (Fig. 3-4), scales (Fig. 4K & Appendix
6
3.2), otolithes (Appendix 4.1), fin spines (Appendix 4.2), fragmented bones, isolated
precaudal & caudal vertebrae, and possible coprolites. They are abundant and
appear frequently, although only the dental components are considered so far to be
sufficiently informative as remains of the fish palaeofauna.
6. Institutional abbreviations
RM PZ, Department of Palaeozoology, Swedish Museum of Natural History,
Stockholm, Sweden; UU, Department of Paleobiology, Uppsala University, Uppsala,
Sweden; LO, Department of Earth and Ecosystem Sciences, Lund University, Lund,
Sweden; PMU, Palaeontological collections, Museum of Evolution, Uppsala
University, Sweden.
Table 1: Occurrences of Upper Cretaceous actinopterygian taxa within the Kristianstad
Basin: (x) = Presence (-) = Absence
Taxa – Localities
Anomoeodus subclavatus
Anomoeodus sp.
Indet. Pycnodontidae
Enchodus cf. E. gladiolus*
cf. Enchodus sp.
Indet. Ichthyodectidae
cf. Pachyrhizodus sp.
Protosphyraena sp.
Indet. Teleost A.
Åsen
x
x
x
x
x
x
Ivö Klack
x
x
-
Ignaberga
x
-
Ullstorp
x
x
x
-
Balsvik
x
x
-
Scania
x
-
*Scania = unknown locality
7. Systematic palaeontology
Class: Osteichthyes Huxley, 1880
Actinopterygii Cope, 1887
Neopterygii Regan, 1923
Order: Pycnodontiformes Berg, 1940
Family: Pycnodontidae Agassiz, 1833 sensu, Nursall 1996
Genus: Anomoeodus Forir, 1887
Anomoeodus subclavatus (Agassiz, 1833)
Fig. 3A-E
Type species: Anomoeodus subclavatus (Agassiz 1833) from Maastrichtian deposits
in the Netherlands.
Referred material: PMU 27255, right prearticular dentition; PMU 27256; left
prearticular dentition; PMU 27257, right prearticular dentition; RM PZ P15753, two
left prearticular dentitions; RM PZ P13746, one isolated prearticular tooth; RM PZ P.
13530, one isolated prearticular tooth; RM PZ P 13743, one isolated prearticular
tooth; RM PZ P13778, one isolated prearticular tooth; LO 11908t, one broken
prearticular tooth.
7
Locality and horizon: Ivö Klack (Blacksudden); Ignaberga (site 1) and Åsen.
(Balsvikensis green), Kristianstad, Scania, Sweden.
Previous designation: Coelodus subclavatus
Description: The most articulated specimens belonging to this taxon consist of
prearticular dentition on white limestone matrix. The specimens exhibit crushing teeth
on the prearticulars. No marginal ornamentation (i.e. grooves or crenulations) can be
recognized on the dentition. The dentition is built up by a main prearticular row with
alternating subcircular to oval shaped teeth sporadically appearing on the posterior
edge (PMU 27256). The main prearticular row exhibits reniform tooth morphology
and the teeth are tightly packed only separated by 1 mm. The prearticulars on the
principal series consists of seven transversely elongated and obliquely placed teeth.
The largest tooth on the main prearticular row in PMU 27256 measures in height up
to 31 mm (anteroposteriorly) and 10 mm in diameter. A decrease in tooth size from
anterior to posterior is evidently shown among the specimens. All tritoral teeth on the
main series, has a concave border facing in an anterior direction. The medial tooth
sequence is partially broken in PMU 27256. Although, the medial teeth have the
same button-shaped to sub-circular morphology and measures in height up to 10 mm
and ~ 0.4 mm in diameter. The medial tooth sequence consists of one row and
appears to be arranged posteriorly. In total, eight medial teeth are recognized in PMU
27256. Moreover, the medial teeth are closely positioned to the principal series. No,
lateral crushing teeth have been persevered in PMU 27256 and only one can be
distinguished in specimen PMU 27255.The teeth shown in RM PZ P 15753 has a “V”
shaped incision on their inner surface indicating wear abrasion.
Discussion: Pycnodontiformes is a monophyletic extinct non-teleostean
actinopterygian group known mainly from Mesozoic deposits (Nursall, 1996a; PoyatoAriza & Wens, 2002; Kriwet, 2005a). As a morphological and ecological distinctive
order of actinopterygians, the pycnodont fishes are mainly recognized by their deep
and laterally compressed bodies (Nursall, 1996a, b; Kriwet, 1999c, 2002a, 2005a)
and also by their durophagous dentition distributed across both the prearticulars and
vomer (Poyato-Ariza, 2003). The paleontological record of pycnodont remains ranges
from the Late Triassic (Norian) to the Eocene (Lutetian-Ypresian) and is represented
by complete skeletons but mostly by isolated dentitions (Kriwet, 2002a; Martín-abad
& Poyato-Ariza, 2013). The neopterygian order Pycnodontiformes include nearly 650
nominal species, belonging to 38 genera, 10 of which are only known from their
dentition and only 78-80 species (about 12 %) are known from their skeletal remains
(Kriwet, 2001b, 2004c, 2005a).
Kriwet (2002a) concluded that Anomoeodus species were a dominant
faunal component during the Upper Cretaceous in Europe. Anomoeodus comprises
nearly 30 species of which only 4 have been recognized on remains aside from
isolated dentition (Kriwet, 1999c; 2002a; Poyato-Ariza & Wens, 2002). The fossil
record of Anomoeodus species has a stratigraphic range from the Late Jurassic
through the Eocene (Shimada & Everhart, 2009) with the best preserved specimens
coming from the Upper Cretaceous of England (Woodward, 1902-1912; Kriwet,
2002a) and other parts of Europe (e.g. Licht & Kogan, 2011).
Two different taxa of Anomoeodus have been recognized (see below)
from the Upper Cretaceous marine deposits of the Kristianstad Basin, the most
common being A. subclavatus. The remains of this taxon are mostly centered and
found at the Ivö Klack locality. Meanwhile, only one isolated as well as broken A.
8
subclavatus tooth has been recovered from the Åsen collection. Specimens of A.
subclavatus have been documented from Maastrichtian deposits in the northeast of
Belgium and the southeast of the Netherlands (Jagt & Dols, 2010; Friedman, 2012).
The identification of A. subclavatus from Upper Cretaceous (Campanian) deposits
from southern Sweden forms a prominent portion of the recovered fish assemblage
and adds additional information to the trophic structure of the Campanian marine
ecosystem of the Kristianstad Basin.
Anomoeodus sp.
Fig. 3F
Referred material: RM PZ P 13670, one tooth; PMU 27261, two isolated
prearticular/vomerine teeth.
Locality and horizon: Balsvik (Stora brottet); Ullstorp, Kristianstad, Scania, Sweden.
Description: In total three specimens has been assigned to this taxon. The material
comprises isolated worn pycnodontiform tooth caps. The specimens exhibits a
concave basal morphology (weakly reniform). The teeth measures in height up to 14
mm and 0.5 mm in diameter and are filled by dentine. The crunching teeth lack root
material and exhibits a smooth occlusal outer surface.
Discussion: As usual with isolated dental remains of pycnodontiform fishes, there are
issues regarding alpha-level taxonomy that must be taken into consideration (Forey
et al., 2003; Buscalioni et al., 2008). Poyato-Ariza (2003) ascertained dental
taxonomy of pycnodont fish as parataxonomic although this was refuted by Kriwet
(2005a).
Subtle characters have been used here to infer taxonomy. Those include:
the hollow nature of the basal aspect, and the concave basal to canoe-like dental
morphology. These characters have been listed as being diagnostic for Anomoeodus
phaseolus by Case & Schwimmer (1988, p. 297-299). Unfortunately, not that many
comparative specimens are available hence leaving the taxonomic designation of this
taxon unresolved. Although, there is no doubt that the teeth discussed here clearly
differ in general morphology from those attributed to Anomoeodus subclavatus. Teeth
of Anomoeodus sp. resemble those of A. phaseolus (see Fowler 1911, p. 147, fig. 90
for comparison).The nominal species Anomoeodus phaseolus is only known from
isolated dentition and has previously only been reported from North America (Fowler,
1911; Case & Schwimmer, 1988).
Indet. Pycnodontidae
Fig. 3G-H
Referred material: LO 11910t, three teeth; LO 11911t, one tooth; PMU 27263, two
teeth; PMU 27260, two vomerine tooth caps.
Locality and horizon: Åsen (Oysterbank & Balsvikensis green); Ullstorp, Kristianstad,
Scania, Sweden.
Description: Teeth of this unknown pycnodontiform taxon are mostly worn, small
sized, measuring in length up to 0.6 mm and 0.5 mm in diameter. In occlusal view the
tooth crown is circular to subcircular in shape and exhibit a smooth punctuated
9
surface. The teeth are filled with dentine and mostly lack root material. In addition the
teeth exhibit slight morphological variation in between each other some being oval in
shape.
Discussion: Dental specimens described herein are recognized as pycnodontiformes
and are infrequently encountered in deposits at Åsen. Ecomorphologically these
teeth are suited for fish with a durophagous diet (e.g. Poyato-Ariza & Wenz, 2002).
This is in agreement with the relatively clear signs of wear that have likely been
produced by mechanical chewing action (e.g. Kriwet, 2001b). Crushing teeth of this
sort are common among the pycnodontiformes. These dental remains have
previously been considered to be abundant in Åsen deposits (Eriksson et al., 2011),
but such degree of abundance cannot be recognized nor supported by the amount of
remains recovered of this taxon in this study.
Figure 3. Teeth of Pycnodontiformes from the Kristianstad Basin. A. Anomoeodus
subclavatus, PMU 27256. B. Anomoeodus subclavatus, PMU 27257. C-D. Anomoeodus
subclavatus, RM PZ P15753. E. Anomoeodus subclavatus, RM PZ P 13743. F. Anomoeodus
sp., RMPZ P 13670. G-H. Indet. Pycnodontidae, PMU 27260 and LO 11911t. Orientations:
A-H, occlusal view. Scale: A–F = 10 mm; G–H = 0.4 mm.
10
Figure 4. Dental remains of stem-group teleosts (Elopiformes, Aulopiformes,
Pachycormiformes, Ichthyodectiformes, and indeterminable teleost fish) from the Kristianstad
Basin. A. cf. Pachyrhizodus sp., LO 11905t. B-C. cf. Pachyrhizodus sp., LO 11907t. D.
Enchodus cf. E. gladiolus, RM PZ P 13685. E. cf. Enchodus sp., RM PZ P 13532. F.
Protosphyraena sp., RM PZ P 13583. G. Protosphyraena sp., PMU 24768. H-I. Indet.
Ichthyodectidae, LO 11909t. J. Indet. Ichthyodectidae, LO 11913t. K. Indet.
Ichthyodectiformes, PMU 27301. L. Indet. Teleost A., LO 11912t. M. Indet. Teleost A., PMU
27262. Orientation: A-M, lingual view; C-I, lateral view; K, anterior view. Scale: A-M = 0.4
mm; K = 10 mm.
11
Subdivision Teleostei Müller, 1845
Order Aulopiformes Rosen, 1973
Suborder Enchodontoidei Berg, 1940
Family Enchodontidae Woodward, 1901
Genus Enchodus Agassiz, 1835
Enchodus cf. E. gladiolus (Cope, 1872b)
Fig. 4D
Cimolichthys gladiolus Cope, 1872, p. 353, 356
Enchodus gladiolus (Cope, 1872)
Type species: Enchodus lewesiensis (Mantell, 1822), from the Upper Cretaceous
(Turonian), Sussex, England.
Referred material: RM PZ P 13685, isolated dental remain of Enchodus cf. E.
gladiolus.
Locality and horizon: (?) Hedvigsdal (Stationen) Ystad, (Scania/Skåne), Sweden. The
locality identification is most likely erroneous.
Description: Specimen RM PZ P 13685 is a palatine tooth of Enchodus cf. E.
gladiolus. The tooth is tall and has a triangular shape. It measures up to ~ 9 mm in
height and ~ 3 mm in diameter. The tooth is laterally compressed and distally curved.
Serrations on the cutting edges are absent and the specimen is slightly sigmoidal in
outline. Longitudinal striations on the midline axis are present at its outer surface and
probably on its inner one as well. In addition, the tooth exhibits both anterior and
posterior cutting edges. The apical portion of the tooth is slightly worn exposing the
inner dentin layer. No post apical barb can be distinguished nor recognized in this
specimen which is a common dental feature seen in E. gladiolus.
Discussion: The greatest issue of concern with the one specimen attributed to
Enchodus cf. E. gladiolus is that the site of discovery is unknown. Here, it is reported
as a part of the Scanian fossil fish collection and not necessarily as a part of the
Kristianstad Basin. Enchodus species have been reported to be very common in
Upper Cretaceous deposits from the Northern Hemisphere (Goody, 1976; Kriwet et
al., 2006). It has been shown that there are five valid species (Robb, 2004; Parris et
al., 2007) belonging to the genus Enchodus in North America: E. ferox Leidy, 1855,
E. petrosus, E. gladiolus Cope, 1874, E. shumardi Leidy, 1856 (cf. Goody, 1976) and
E. dirus Leidy, 1857b. Enchodus remains have also been reported from Lebanon
(Forey et al., 2003), Palestine and the Negev (Chalifa & Lewy, 1991; Chalifa, 1996),
Syria (Bardet et al., 2000), Saudi Arabia (Kear et al., 2009), North Africa
(Arambourg, 1952), Netherlands (Goody, 1968), Belgium (Leriche, 1929; Friedman,
2012), England (Longbottom & Pattersson, 1987), Antarctica (Kriwet et al., 2006),
Czech Republic (Wiese et al., 2004), Brazil (Reboucas & Da Silva Santos, 1956) and
several other countries, proving the worldwide distribution of Enchodus during the
Cretaceous.
Enchodus being a pelagic mid-food chain predatory fish is considered to
have been able to inhabit a wide range of ecological niches (Fishman et al., 1995;
Fielitz, 2004). The occurrence of Enchodus gladiolus has been documented from a
variety of localities in the Upper Cretaceous of North America (Becker et al., 2009).
Enchodus is known from Early Cretaceous deposits (Albian) to the Palaeocene
12
(Winkler et al., 1990; Forey et al., 2003) meanwhile the stratigraphic range of
Enchodus gladiolus begins in the Turonian and extends into the Late Maastrichtian
(Goody, 1976; Fielitz, 1996; Robb, 2004; Shimada et al., 2006). The Palaeocene
occurrence of Enchodus reported by Arambourg (1952) has been questioned (Cavin,
2001) and consequently been regarded as being restricted to the Cretaceous.
The identification of specimen RM PZ P 13685 has been in accordance
with the diagnosis given by Goody (1976, p. 103-104) and also with numerous
figured comparative specimens (e.g. Shimada et al., 2006, p. 25-26, fig, 14.1-14.5;
Nagrodski et al., 2012, p. 8-9, fig, 4L; Becker et al., 2009, p. 3-5, fig, 2I; Jansen et al.,
2013, p. 5, fig, 3H; Cumbaa et al., 2010, p. 208, fig, 5T; and others).
cf. Enchodus sp.
Fig. 4E
Referred material: RM PZ P 13532, one tooth; LO 11906t, one tooth.
Locality and horizon: Balsvik (Norra brottet); Åsen (Balsvikensis yellow), Kristianstad,
Scania, Sweden.
Description: The material consists of two isolated, presumably palatine teeth of cf.
Enchodus sp. The teeth measures in height up to 0.5 mm and has a diameter length
of 0.2 mm. The teeth have a smooth un-striated outer surface and unserrated cutting
edges. The teeth are slightly sigmoidal in outline and possess a clear and wide open
pulp cavity at the base of the tooth. Specimen RM PZ P 13532 is partially broken
along its right outer margin exposing the inner dentine layer. In addition, the teeth are
covered by shining smooth enamel.
Discussion: The recovered specimens suggest the occurrence of more than one
Enchodus taxon which coexisted at the time of the Upper Cretaceous of southern
Sweden. Teeth belonging to cf. Enchodus sp. are small and ecomorphologically
characteristic of fish with an ichtyophagous diet. Enchodus species have previously
been interpreted as being open water predatory fish, feeding on fast swimming
cephalopods (Grandstaff & Parris, 1990).Fossil remains of Enchodus are not
abundant and currently only represented by teeth making them extremely rare.
Order Ichthyodectiformes Bardack and Sprinkle, 1969
Family Ichthyodectidae Crook, 1892
Indet. Ichthyodectidae
Fig. 4H-K
Referred material: LO 11909t, one presumably palatine tooth; LO 11913t, one
partially broken tooth; PMU 27301, cycloid scale.
Locality and horizon: Åsen (Oysterbank and Balsvikensis green); Bäckaskog,
Kristianstad, Scania, Sweden.
Description: Teeth belonging to this unidentified Ichthyodectiform taxon are nearly
straight and show no significant inclination towards any direction. To note, in lateral
view specimen LO 11909t is slightly curved posteriorly and not sigmoidal. The teeth
are measured in height up to 10 mm and 0.4 mm in diameter. The teeth bear mesial
13
and distal carinae. Both cutting edges are not serrated, although a wavy mesoscopic
pattern can be followed along both margins which extend to the base of the teeth.
The base of the teeth has no distinct pulp cavity and the teeth are covered by a thin
slightly damaged enameloid cover. Both lingual and labial faces presumably bear
vertical striae as it faintly appears to present in specimen LO 11913t.
Discussion: Ichthyodectiform fishes are known from several Mesozoic sites around
the world (e.g. Alvarado-Ortega, 2004; Kriwet et al., 2006; Shimada et al., 2006;
Friedman, 2012; Kim et al., 2014). These ranges stratigraphically from the
Kimmeridgian into the Maastrichtian (Patterson, 1993; Schwimmer et al., 1997b).The
Ichthyodectiformes were common as well as abundant in regions of the Northern
Hemisphere during the Upper Cretaceous (Kriwet et al., 2006).Teeth attributed to
Indet. Ichthyodectidae are lanceolate, slender, tall as well as triangular in outline,
making them distinctive and readily identifiable as Ichthyodectiformes. Here, they
have been recovered from Åsen and are so far only represented by isolated dental
remains. The taxonomic assessment of teeth attributed to Indet. Ichthyodectidae is
sufficient for now, pending for the acquisition of taxon specific elements.
Order Pachycormiformes Berg, 1940
Family Pachycormidae Woodward, 1895
Genus Protosphyraena Leidy, 1857a
Protosphyraena sp.
Fig. 4F-G
Type species: Erisichthe nitida Cope, 1872c, from Upper Cretaceous strata of the
Niobrara Chalk, Kansas, USA.
Referred material: RM PZ P 13583, one tooth; PMU 27258, one tooth; PMU 27259,
one tooth; PMU 24768, one tooth.
Locality and horizon: Ivö Klack (Site 1) Ivö; Ullstorp, Kristianstad, Scania, Sweden.
Description: Isolated dental remains of pachycormid fish are rare finds within the
Kristianstad Basin, yet their distinctive characteristics make them readily identifiable.
The dental (19 mm high and 8 mm wide in RM PZ P 13583: Fig. 4F) remains are
represented by a few isolated tooth crowns. The general morphology shared among
the specimens comprises a large-sized, arrowhead shaped, labio-lingually
transversally compressed tooth. Moreover, the teeth exhibit longitudinal post mortem
cracks on their inner and outer surfaces and bear well-defined anterior and posterior
carinae. Both cutting edges are unserrated and slightly convex along both margins.
Tooth apices show sign of breakage and wear in all specimens exposing the inner
dentin layer.
Discussion: Upper Cretaceous (Campanian – Maastrichtian) pachycormid remains
are currently represented only by the comparable swordfish-like Protosphyraena
(Stewart, 1988a; Lambers, 1992). This taxon has been reported from various
countries including: England (Woodward, 1902-12), North America (Bardack, 1968;
Applegate, 1970; Stewart, 1988a; Everhart, 2005; Shimada et al., 2006), Belgium
(Leriche, 1929; Friedman, 2012), Spain (Vullo et al., 2009), Netherlands (Diedrich,
2001), Australia (McNamarra et al., 1993) and Saudi Arabia (Kear et al., 2009).
14
Dental remains being so far the only skeletal component recovered from this Upper
Cretaceous teleost fish verily represent their first encounter in the Kristianstad Basin.
However, the remains of Protosphyraena are scarce in deposits from the Kristianstad
Basin. Despite the few recovered dental remains of this stem teleost group it has
been proven to be enough to conclude their presence and importance in this former
inhabited marine ecosystem.
Teeth attributed to Protosphyraena has previously been mislabeled and
assumed to be remains of large-sized carnivores dinosaurs (Persson, 1959: p. 474,
pl. 20, fig. 4). This was initial shown by Lindgren & Siverson (2002), where they
concluded that the three dinosaurian dental remains from the Ivö Klack locality,
reported by Persson (1959) were in fact incorrectly labeled and concluded to be
teleostean dental remains. Lindgren et al., (2008) proposed correctly the true
taxonomic affinity of the dental remains and placed them as pachycormids.
Order Elopiformes Greenwood, Rosen, Weitzman, and Meyers, 1966
Family Pachyrhizodontidae Cope, 1872b
Genus Pachyrhizodus Dixon, 1850
cf. Pachyrhizodus sp.
Fig. 4A-C
Referred material: LO 11905t, two teeth; LO 11907t, one tooth; LO 11914t, one tooth.
Locality and horizon: Åsen (Coquina Bed; Oysterbank and Balsvikensis green),
Kristianstad, Scania, Sweden.
Description: Teeth belonging to this genus, measures in height up to 0.6 mm and 0.3
mm in diameter. The teeth possess longitudinal oriented striations on their conical
shaped crown. The covering of the tip of the teeth (i.e. the crown) is made up of a
thin yet shining enameloid cap. The teeth have two well defined cutting edges
although they are not serrated. The teeth are slightly inclined lingually and possess a
distinct clear pulp cavity at the base and are subcircular in cross section.
Discussion: The true taxonomic identity of teeth attributed to cf. Pachyrhizodus sp. is
uncertain, but the specimens are referred to cf. Pachyrhizodus sp. due to their
distinctive dental morphological appearance which comprises an intermediate sized,
slightly flexed conical form. These teeth could be the same as described as Indet.
Teleost A, (see below). The teeth of Pachyrhizodus have usually a characteristic
swollen base but this has not been persevered in any of the specimens.
Pachyrhizodontoid are an extinct group of Cretaceous teleosts,
containing fusiform predatory fish such as Pachyrhizodus (Friedman, 2012). The
genus Pachyrhizodus ranges stratigraphically from the Albian of Australia and
England (Bardack, 1962; Bartholomai, 1969; Forey, 1977) to the Maastrichtian of
Belgium, Netherlands and North America (Dollo, 1892; Stewart, 1898a, 1899b;
David, 1946; Applegate, 1970). Teeth of Pachyrhizodus are not abundant in the
samples collected from Åsen and are often damaged. The occurrence of
pachyrhizodontoids at Åsen is particularly interesting because skeletal elements
belonging to Enchodus have previously been recorded in gastric residues of
pachyrhizodontoids (Shimada & Everhart, 2003).
15
Indet Teleost A.
Fig. 4L-M
Referred material: PMU 27262, six teeth; LO 11912t, five teeth.
Locality and horizon: Åsen (Oysterbank), Kristianstad, Scania, Sweden.
Description: Teeth of this unknown taxon measure up to 0.6 mm in height and have a
diameter length of 0.2 mm. The teeth are small sized, triangular in shape and slightly
inclined lingually. Among the specimens only few teeth bear a thin remnant enamel
covering. Most of the teeth exhibit a clear open pulp cavity at the base. These teeth
lack serrations along both cutting edges and striations along the midline axis.
Discussion: The teeth described here are of uncertain systematic affinity but could be
the same as those described as cf. Pachyrhizodus sp. Their shape and form is very
much the same as for those attributed to Pachyrhizodus. But, vital characteristics
such as vertically oriented striations on the conical shaped crown and the presence a
thin enameloid cap at the tip of the tooth are absent in these specimens. Remains of
this unknown taxon are only recovered from the B. mammillatus biozone (Fig. 1) and
comprise a number of isolated partially broken teeth.
8. Discussion
8.1 Palaeoecology of the actinopterygian assemblage
The actinopterygian assemblage of the Kristianstad Basin contains nektonic
durophagous carnivores (Anomoeodus subclavatus, Anomoeodus sp. and the small
sized pycnodont fish attributed to Indet. Pycnodontidae), and piscivoroues forms
(Protosphyraena sp., Indet. Ichthyodectidae, cf. Pachyrhizodus sp., Teleost Indet. A,
and with some uncertainty Enchodus cf. E. gladiolus). Remains of Anomoeodus
subclavatus have been recovered from Ivö Klack, Ignaberga and Åsen while
specimens belonging to Anomoeodus sp. have been recovered from Ullstorp and
Balsvik. Dental remains attributed to Protosphyraena sp. are found at the Ivö Klack
and Ullstorp localities. Bony fish specimens recovered at Åsen belonging to Indet.
Ichthyodectidae and cf. Enchodus sp. are few in number, but the occurrence of cf.
Enchodus sp. from Balsvik concludes the presence of Enchodus species in more
than one locality within the Kristianstad Basin. As for the specimens that have been
attributed to cf. Pachyrhizodus sp. they are so far only known from the Åsen material
and the same applies for specimens attributed to Indet. Teleost A.
Despite the fact that pycnodont fish have been recovered from most of
the Kristianstad Basin localities, it is premature to suggest that the actinopterygian
fauna maintained a similar taxonomic composition across the entire basin. Based on
what has been recovered so far, it is plausible too suggest that future studies will
show slight differences in the taxonomic composition of actinopterygians across the
basin.
All actinopterygian taxa reported herein represent the first record of
Upper Cretaceous bony fish documented from the Fennoscandian shield. The
abundance and pattern of diversity of Upper Cretaceous actinopterygian fish from
southern Sweden, is yet not fully understood and requires further investigations. That
being said, all five bony fish families from the basin had during the Cretaceous a
worldwide distribution and were important faunal components in former marine
16
palaeocommunities (e.g. Nicholls & Russell, 1990; Shimada et al., 2006; Kriwet et al.,
2006; Kear et al., 2009; Nagrodski et al., 2012).
Dental remains of pycnodontiforms are the most common identifiable
actinopterygian remains hitherto recovered from the Kristianstad Basin. These are
followed by indeterminable teleost teeth and vertebrae, and by dental remains
attributed to Protosphyraena, Pachyrhizodus, Enchodus and indeterminable
ichthyodectiform fish. The sample size of bony fish that has been received from the
Ignaberga, Ullstorp and Balsvik localities is very small and comprises only few dental
remains, unlike the Åsen material that comprises hundreds if not thousands of bony
fish elements.
The relative abundance of carnivorous actinopterygian fishes reported
from the Kristianstad Basin together with previously reported elasmobranch and
aquatic amniotes (Sørensen et al., 2013) were part of a palaeocommunity diverse in
organisms found at the lower trophic levels (Sørensen et al., 2013). Siverson (1992a)
even suggested that the diverse selachian fauna uncovered at the Åsen site may be
explained solely on the basis of the great abundance of belemnites.
The Campanian fish fauna from the Kristianstad Basin is
palaeobiogeographically similar to Boreal assemblages, suggesting faunal
communications (e.g. the English fish fauna) (see Longbottom & Pattersson, 1987).
This is further supported by faunal similarities shared between the North American
Western Interior Seaway and the Kristianstad Basin which is moderately high both on
the generic and species level. Such similarities were briefly mentioned by Rees &
Lindgren (2005) dealing with the hesperornithiformes genus Baptornis which turned
out to be their first record outside the Western Interior Seaway. Lindgren & Siverson
(2004) documented and demonstrated both the occurrence of the mosasaurine taxon
Clidastes propython in southern Sweden and the paleobiogeographical extension of
the species beyond North America. All reported actinopterygian taxa from southern
Sweden are known from Upper Cretaceous deposits in the Western Interior Seaway
(e.g. Nicholls & Russell, 1990; Becker et al., 2009; Cumbaa et al., 2010; Nagrodski et
al., 2012). The degree of similarity shared in between several Upper Cretaceous
fossil sites around the world, in particular the fishes; suggest that passages were
utilized by cosmopolitan fish throughout the North American Seaway, the Boreal and
Tethyan realms (Cumbaa et al., 2010).
All present data so far suggests that the bony fish fauna from the
Kristianstad Basin is far from being complete. This is strongly supported by simple
comparisons with contemporaneous faunal assemblages across the world and in
particular with the Western Interior of North America.
Based on the amount of recovered bony fish elements from the Åsen
locality (Fig. 6), a diverse actinopterygian/teleostean fauna with variable feeding
ranges is indicated to have been present. The preservation status of taxonomically
identifiable bony fish remains is poor but further field recovery will likely add
additional actinopterygian taxa to the faunal list.
8.2 Pycnodontiformes
The pycnodontiformes were key faunal members of most Mesozoic fish assemblages
globally (Martín-abad & Poyato-Ariza, 2013) and their presence and diversity during
the Upper Cretaceous of the Kristianstad Basin warrants biogeographical
comparisons. Dental remains of pycnodont fish have been recovered from the marine
deposits at the Åsen site. These are currently not fully identified and comprise only
17
few isolated vomerine tooth caps. Also from Ivö Klack and Ullstorp, several
generically indeterminable specimens of pycnodont character have been
encountered, as such suggesting their relative abundance and diversity in the
Kristianstad Basin. These are represented by disarticulated specimens comprising
stout, crushing teeth reflecting their grazing durophagous diet. Also, branchial teeth
of pycnodont fish are previously known from the B. mammillatus biozone (M.
Siverson, pers.com.) and newly recovered (not described) non-teleostean specimens
from Ullstorp (Appendix 3.1) may be assigned to aff. Coelodus sp.
Pycnodont fish inhabited a wide variety of environmental settings, but are
mostly encountered in coastal marine deposits (Nursall, 1996a; Martín-abad &
Poyato-Ariza, 2013). The record of pycnodontiformes points to an origin around the
Western Tethys (Martín-abad & Poyato-Ariza, 2013).
The great abundance of shelled invertebrate groups at Ivö Klack, Ullstorp
and Åsen, which are known to have been exploited by pycnodont fish (Nursall,
1996b) (e.g. reefal organisms such as bivalves and echinoderms), may had served
as a vital resource for the pycnodont fishes and in addition it may explain the
abundance of pycnodont fish in deposits at Ivö Klack, Ullstorp and with some
uncertainty at Åsen as well (see discussion above). To note, teeth of pycnodont fish
have recently been recovered from the gravel pit at Bäckaskog, Scania, Sweden (T.
Engfors, pers.com.).
8.3 Aulopiformes
The Enchodontid Enchodus represent an extinct group of fishes known to have had a
worldwide distribution during the Upper Cretaceous (Chalifa, 1996). These are placed
within the alepisauroid lancetfishes (Kriwet, 2003) and possess characteristic palatine
fang-like teeth (Longbottom & Pattersson, 1987; Fielitz, 1999). Enchodus remains in
the Upper Cretaceous of the Kristianstad Basin are only represented by a few
isolated teeth. Enchodus species were marine pelagic piscivoroues teleost but could
also venture into shallow water environments (Goody, 1976).
Their presence in the Northern Hemisphere is common (Kriwet et al.,
2006) and here they are reported from Åsen and Balsvik. The one specimen that has
been attributed to Enchodus cf. E. gladiolus in the fossil fish collection from RM PZ is
currently quite problematic because it is not known from which locality it derives. In
addition, such dental remains have not been encountered so far from any other fossil
site within the Kristianstad Basin. Despite this, it is premature to suggest that the one
tooth of Enchodus cf. E. gladiolus is not from any locality in Sweden. Also, if correctly
identified the one tooth of E. gladiolus would represent the first record outside North
America.
8.4 Ichthyodectiformes
The identification of specimens attributed to Indet. Ichthyodectidae are represented
by two isolated palatine teeth. Morphologically consistent dental remains from the
upper Campanian-? early Maastrichtian Herbert Sound Member in Antarctica were
figured and identified by Kriwet et al., (2006, fig. 4i) as indeterminable
Ichthyodectiformes. These are tall, laterally compressed, unserrated along both
margins, slender and lanceolate. In addition, the Åsen specimens (LO 11909t and LO
11913t, Fig. 3H-J) attributed to the Ichthyodectiformes appear to be compatible with
those of Gillicus from the Upper Cretaceous of North America. They differ from the
18
North American Xiphactinus in both size and morphology (e.g. Shimada et al., 2006:
fig. 13; Schwimmer et al., 1997b: fig. 2A-C) and are larger than those found in
Ichthyodectes (e.g. Bardack, 1965).
Associated cycloid scales from Bäckaskog (Fig. 4K) may positively be
assigned to indeterminable Ichthyodectiformes (see Friedman, 2012: fig. 2B; Kriwet
et al., 2006: fig. 4E-G, for comparison).These are dorsoventrally ovoid to circular in
outline and wider than long. The most complete scale is marked by numerous
concentric circuli and several distinct anterior radii that extend from the oval shaped
focus. Additional ornamentations such as punctae on the inner part and/or on the
posterior edge cannot be seen. These are large sized, (0.9 mm high and 14 mm wide
in PMU 27301: Fig. 4K) preserving several features but unfortunately of low
taxonomic value and here allocated to indet. Ichthyodectiformes.
8.5 Pachycormiformes
The Kristianstad Basin pachycormid remains comprises so far only few and often
damaged isolated teeth. Such dental remains are morphologically similar to those of
the Upper Cretaceous pachycormid genus Protosphyraena Leidy, 1857 (see
Shimada et al., 2006: fig. 13, 2-3; Nagrodski et al., 2012: fig. 4C for comparison).
These dental remains are often of large-sized suggesting a substantially sized
predator. The protosphyraenids were likely among the largest bony fish to have lived
in the shallow waters of the relatively small sized basin at the time of the Upper
Cretaceous. Being large-sized, swift and predaceous teleost fish (Schwimmer et al.,
1997a; Arratia, 2004), the presence of Protosphyraena in the Upper Cretaceous
deposits of southern Sweden, together with previously reported elasmobranch
species and marine amniotes, attests to the richness and diversity of predaceous
organisms at the Kristianstad Basin during the Upper Cretaceous.
8.6 Elopiformes
The genus Pachyrhizodus had a worldwide distribution in the Cretaceous (e.g.
Romer, 1966; Wiffen, 1983; Cumbaa et al., 2010). Teeth attributed to Pachyrhizodus
from the Kristianstad Basin are few, isolated and lack an expanded base. The
Pachyrhizodontidae comprised piscivoroues fishes and were most diverse in regions
of the Western Hemisphere (Cavin, 2001). The amount of recovered dental remains
attributable to Pachyrhizodus at Åsen suggests that they presumably were an
important faunal component of the Upper Cretaceous palaeocommunity.
9. Taphonomic and palaeoenvironmental implications
The Campanian vertebrate assemblage of the Kristianstad Basin was dominated by
aquatic forms such as selachian and marine reptiles (Siverson, 1992a, b; Lindgren,
2004, 2005). Also few continental forms including ornithischian and recently
recovered dental remains of theropod dinosaurs have been recorded in the Upper
Cretaceous of the Kristianstad Basin (Lindgren et al., 2007; E. Einarsson et al.,
unpublished data). The Åsen site has previously been interpreted as a coastal
environment surrounded by shallow murky waters (Rees, 1999). The nature of
preservation of bony fish elements as well as other fossil remains attests to a high
energy environment. A low to intermediate sedimentation rate at the Åsen site may
be proposed based on the degree of disarticulation as well as fragmentation of
19
shelled organisms, in particular bivalves. Although articulated oysters, are frequently
encountered in the upper section of the B. mammillatus biozone (Lindgren &
Siverson, 2004). A taphonomic bias in terms of preservation is proposed here since
the recovered sample of bony fish comprises only disarticulated specimens all
showing signs of abrasion. This is further supported by the prevalence of resistant
elements such as teeth (Fig. 6) which may indicate extensive post-mortem
mechanical destruction. The concentrate comprises altogether > 10 000 fish bone
fragments, vertebral central, dental remains of sharks, teeth, vertebrae and bones of
squmates, turtle bones, mandibular and additional skeletal components of
chimaeroids, bivalvian shells and other invertebrate remains and possible plant
material.
Actinopterygian
Chondrichthyan
NUMBER OF TAXA
6
5
4
3
2
1
0
FAMILIES
Figure 5. Diagrammatic representation of the total amount of documented fish families and
the number of taxa within each taxonomic group, present in the Kristianstad Basin. Based
on data compiled by Sørensen et al., (2013: table 2) and this paper.
More than sixty elasmobranch species have previously been recorded in
the Cretaceous deposits of the Kristianstad Basin of which nearly forty cartilaginous
fishes are known from Åsen (Siverson, 1993b). Among the most frequently
encountered are lamniform sharks including Archaeolamna kopingensis, Carcharias
latus, and Squalicorax kaupi (Siverson, 1992a, 1993b). The chondrichthyans are
usually represented by dental remains which attest to their resistance and durability
during pre-burial transport. They are abundant (i.e. the teeth) and simply reflect the
chondrichthyans ability to replace a huge amount of teeth during their lifespan. This is
in contrast with the actinopterygians whose dental remains have a much lower
resistibility towards mechanical destruction, which is supported by the low
preservation of dental remains. Most resisting towards post mortem mechanical
destruction among the actinopterygians is teeth attributed to indeterminable teleost
fish which make up 44 % of all actinopterygian dental remains at the Åsen site.
Despite the taphonomic losses of identifiable actinopterygian remains at Åsen, recent
reports published on the vertebrate fauna of the Kristianstad Basin (e.g. Siverson,
1992a; Lindgren & Siverson, 2002, 2004; Lindgren et al., 2007; Einarsson et al.,
2010; Sørensen et al., 2013) highlights a collecting and research bias towards larger
20
vertebrates, resulting in an under-representation of the actinopterygian fauna. This
does not mean that the actinopterygian fauna was less diverse, than any other
vertebrate group rather it reflects taphonomic losses as well as collection biases.
Also, the overwhelming amount of recovered teleostean vertebrae and additional
skeletal elements from Åsen suggest that plenty of fish species still remain
unidentified, hence concluding that the actinopterygian fauna was both ecologically
and taxonomically diverse. In addition, trophic transfer efficiency analyses of the
Kristianstad Basin based on the presence of a diverse assemblage of larger apex
predators suggest that sufficient prey items (e.g. smaller fish and benthic
invertebrates) must had been present in turn to sustain the ecosystem (Sørensen et
al., 2013).
The marine strata at the Ivö Klack site were deposited within the intertidal
zone of a rocky coastal environment with shallow waters (Surlyk & Christensen,
1974). The bony fossil fish specimens that derive from the Ivö Klack site are better
preserved than those examined at the Åsen site. Essentially all fossil vertebrate
elements uncovered at Ivö Klack tend to be poorly preserved and are frequently
encountered as isolated bone and teeth (Surlyk & Sørensen, 2010). This is true for all
Cretaceous fossil sites within the Kristianstad Basin and as pointed out by Lindgren &
Siverson (2002) it simply reflect environmental parameters including a near-shore,
high energy depositional setting with intense currents (i.e. reworking and
transportation) and the presence of many scavenging organisms (i.e. resulting in
disaggregation and dispersal). Still the role played by sediment type (i.e. carbonates
at Ivö Klack and siliciclastic marine strata at Åsen) may affect the final preservation of
bony fish components.
Bulk sample from Åsen
907
∑
Total specimens
1000
85 % shark
15 % bony fish
378
800
259
600
169
400
200
17
55
2
31
16
7
0
1
0
Coquina Bed Oysterbank
Actinopterygian teeth
Balsvikensis
green
Chondrichthyan teeth
Actinopterygian vertebrae
Actinopterygian teeth
Balsvikensis
yellow
Actinopterygian vertebrae
Chondrichthyan teeth
Figure 6. Quantitative composition of actinopterygian remains in stratigraphic order at the
Åsen locality. Bony splinters, fin spines and additional skeletal components of
actinopterygian fish are not considered in the diagram. Although they make up, a dominant
portion of the total bony fish biomass.
Based on the degree of disarticulation of all major vertebrate fossil
groups at the Åsen site, the fossil assemblage is interpreted as an allochthonous
21
taphocoenoses. This is further supported by the lower B. mammillatus coquina bed
which represents a storm generated deposit with a moderate degree of transportation
(Lindgren & Siverson, 2002). The degree of transportation of fossils forming the
coquina bed is not high as pointed out by Eriksson et al., (2011) and may instead be
explained by hydrodynamic reworking (Eriksson et al., 2011). The abundance of
gastric residues found at Åsen are considered to have undergone rapid burial and
lithification followed by taphonomic processes involving transportation and reburial
prior to storm events (Eriksson et al., 2011). In either case, the fossil assemblage at
Åsen has been subject to complex taphonomic processes, which is supported by the
presence of a lag deposit overlying the B. mammillatus biozone which presumably is
associated with sea level cyclicity consequently resulting in reburial of debris and
associated fossil contents (e.g. Becker et al., 2004, 2006b, 2009; Shimada et al.,
2006). Such environmental conditions are not conducive to preservation of poorly
ossified skeletal elements of bony fish (e.g. Becker et al., 2009).
Sea level fluctuations are recorded in the stratigraphic sequence at Åsen,
with the presence of glauconitic greensand strata and a high biodiversity in the B.
mammillatus biozone indicating transgression. The B. balsvikensis biozone is in
general highly unproductive in its fossil content in comparison with the B.
mammillatus biozone containing mostly benthic fossil organisms such as bryozoans
and sea urchins. The faunal composition of actinopterygian fish at Åsen show slight
differences throughout the stratigraphic succession (Fig. 1). Pycnodont teeth being
so far documented from both the B. mammillatus and B. balsvikensis biozones
(Fig.1) are mostly concentrated in the oysterbank and appear occasionally in the
balsvikensis green horizon. The Ichthyodectiformes being less abundant however
maintain the same stratigraphic range as those attributed to the pycnodontiformes.
The appearance of dental remains of Pachyrhizodus begins at the coquina bed and
disappears in the top stratigraphic section (i.e. balsvikensis yellow). As for the
enchodontids they have their first and last appearance in the balsvikensis yellow
horizon. The uppermost horizon ( 1.6 m/thick) at Åsen has produced only a few fish
remains which contrasts all underlying layers especially the oysterbank (0.6 m/thick)
which have produced more fish elements (Fig. 6).
A major faunal turnover at the lower/upper Campanian boundary has
been proposed by Lindgren & Siverson (2004) based on the compositional
differences between the B. mammillatus and B. balsvikensis biozones. The few
recovered actinopterygian specimens from the B. balsvikensis biozone (Fig. 6) attest
to the fact that major perturbations must have taken place. The lithological
differences between the two local biozones at Åsen represent a shift in facies and
may serve as an explanation to the major drop in overall biodiversity in the upper
stratigraphic sequence. Lindgren & Siverson (2004) have argued for an
intercontinental isochronous event and/or a sequence of events which caused a
major decline in mosasaur communities both in North America and the Kristianstad
Basin. The argument put forward by Lindgren & Siverson (2004) being valid is
however not conclusive as to the fact that not only does the mosasaur fauna
disappear at the lower/upper Campanian boundary at Åsen, but so does most other
marine vertebrates as well. As to whether the biotic turnover is real or an artifact of
taphonomy require further investigations.
22
Numerical teeth count
40
35
30
25
20
15
10
5
0
34
11
3
PYC
ENC
2
4
ICH
PAC
Actinopterygians
Bulk sample
4
PAY
Indet T
Figure 7. Quantitative composition of the actinopterygian fauna from the Kristianstad basin,
Scania, Southern Sweden (number of teeth per family collected from the samples). PYC =
Pycnodontidae; ENC = Enchodontidae; ICH = Ichthyodectidae; PAC = Pachycormidae; PAY
= Pachyrhizodontidae; Indet T = Indeterminable Teleost (see Appendix 1.1 – 1.3).
10. Conclusions
The Campanian actinopterygian assemblage of the Kristianstad Basin has been
studied for the first time. The present study has elucidated the ecological importance
of the assemblage and its environmental implications. The following concluding
remarks and results are underlined:







The fossil vertebrates encountered in Upper Cretaceous deposits in the
Kristianstad area represent a microvertebrate assemblage.
The vertebrate fauna consists of sharks, rays, rat fishes, bony fishes, marine
reptiles such as mosasaurs and plesiosaurs, sea turtles, aquatic
hesperornithid birds, crocodiles and dinosaurs.
The recovered bony fish specimens are few and often not diagnostic. This
suggests major taphonomic losses which have made it difficult to assess the
fossils taxonomically.
The actinopterygian assemblage of the Kristianstad Basin bears resemblance
towards other Upper Cretaceous faunal assemblages across the world,
suggesting biogeographical communications.
Despite the few identifiable bony fish remains recovered from Åsen, the
abundance of skeletal components such as fin spines and fragmented cranial
and postcranial skeletal bones of actinopterygians indicate that they
presumably formed a prominent portion of the now extinct ecosystem.
The great absence of a presumably diverse teleost fish fauna at Åsen has
opened a taphonomic window to the ecology of this palaeoenvirnoment.
The actinopterygian assemblage of the Kristianstad Basin is ecologically
diverse. It comprises nektonic/nektobenthic durophagous carnivores (e.g.
Anomoeodus subclavatus) and epipelagic piscivoroues forms (e.g.
Protosphyraena sp.).
23


The lower/upper Campanian boundary at Åsen marks the transition from a
healthy and mature marine biotic community to a near absence in marine
vertebrates. Lithological and ecological data suggest local environmental
changes rather than an intercontinental extinction event.
In this paper a new actinopterygian fauna have been described representing
the first record of Upper Cretaceous bony fish from the Fennoscandian shield.
11. Acknowledgments
This work was carried out at the Museum of Evolution, Uppsala University, Sweden. I
extend my most sincere gratitude’s to the following people: Dr. Jan Ove Ebbestad for
teaching me how to use CorelDraw and for his constant support and advice in most
aspects related to this project. I thank my supervisors Dr. Benjamin Kear for financial
support, assistance with image production, and for providing helpful comments on
previous drafts; and Elisabeth Einarsson for her constant support, advices and most
importantly for her patience and willingness to assist me whenever I got stuck. I thank
field paleontologist Tord Engfors for providing me with all fossil specimens from the
Ullstorp locality. Without his contributions to this work, much would have remained
unknown. I thank Dr. Michael Streng for helpful comments on the stratigraphic chart
(Figure. 1) and Professor Graham Budd for providing literature on the nature of the
fossil record. I thank Dr. Mikael Siverson for helpful discussions concerning the
selachian fauna of the Kristianstad Basin. I thank Dr. Henning Blom for providing
literature on the taxonomic utility of otolithes in paleoichthyology. I thank the Åsen
excavation team for sharing their joy in doing paleontology and finally I thank my
good friend Filip Andersson for providing instructions in how to use Adobe
Photoshop.
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32
13. Appendices
Appendix 1
LO
Site and Biozone
LO.
11905t
LO.
11906t
LO.
11907t
LO.
11908t
LO.
11909t
LO.
11910t
LO.
11911t
LO.
11912t
LO.
11913t
LO.
11914t
LO.
11915t
Åsen (Coquina Bed), Bromölla,
Sweden
Åsen (Balsvikensis yellow), Bromölla,
Sweden
Åsen (Balsvikensis green), Bromölla,
Sweden
Åsen (Balsvikensis green), Bromölla,
Sweden
Åsen (Balsvikensis green), Bromölla,
Sweden
Åsen (Balsvikensis green), Bromölla,
Sweden
Åsen (Balsvikensis green), Bromölla,
Sweden
Åsen (Oyster bank), Bromölla,
Sweden
Åsen (Oyster bank), Bromölla,
Sweden
Åsen (Oyster bank), Bromölla,
Sweden
Åsen (Coquina Bed), Bromölla,
Sweden
Identification
∑
cf. Pachyrhizodus sp.
2
cf. Enchodus sp.
1
cf. Pachyrhizodus sp.
1
Anomoeodus subclavatus
1
Indet. Ichthyodectidae
1
Indet. Pycnodontidae
3
Indet. Pycnodontidae
1
Indet. Teleost A.
5
Indet. Ichthyodectidae
1
cf. Pachyrhizodus sp.
1
Teleostei incertae sedis
(Saccular otolith)
1
Appendix 1.1: Registered fossil specimens from the Åsen collection.
33
RMPZ
Site and Biozone
P.15753
P.13530
P.13746
Blaksudden (site 1), Ivö, Sweden.
Blaksudden (site 1), Ivö, Sweden.
Blaksudden (Senon, Mammillatus
Zone), Ivö, Skåne, Sweden.
Blaksudden (Senon, Mammillatus
Zone), Ivö, Skåne, Sweden.
Balsvik (Stora brottet),
Kristianstad, Skåne, Sweden.
Sweden (site 3) Sweden.
(unknown locality)
P.13745
P.13670
P.1416
P.13743
P.13528
P.13778
P.13697
P.13770
P.13698
P.13532
P.13685
R.1437
R.1438
R.1186
R.1298
R.1254
R.1125
P.1386
P.1387
R.1237
R.2217
P.8697
P.13702
P.13583
Blaksudden (Senon, Mammillatus
Zone), Ivö, Skåne, Sweden.
Sweden (Site 3) Sweden.
Ignaberga (site 1) Hässleholm,
Skåne, Sweden.
Blaksudden (Senon, Mammillatus
Zone), Ivö, Skåne, Sweden.
Axeltorp (site 2) Bromölla, Skåne,
Sweden.
Blaksudden (Senon, Mammillatus
Zone), Ivö, Skåne, Sweden.
Balsvik (Norra brottet),
Kristianstad, Skåne, Sweden.
Hedvigsdal (Stationen), Ystad,
Scania/Skåne, Sweden (locality is
most likely wrong).
Blaksudden (site 1), Ivö, Sweden.
Blaksudden (site 1), Ivö, Sweden.
Blaksudden (site 1), Ivö, Sweden.
Blaksudden (site 1), Ivö, Sweden.
Blaksudden (site 1), Ivö, Sweden.
Blaksudden (site 1), Ivö, Sweden.
Annetorp (site 2), Malmö, Skåne,
Sweden (locality is wrong).
Annetorp (site 2), Malmö, Skåne,
Sweden (locality is wrong).
Blaksudden (site 1), Ivö, Sweden.
Blaksudden (site 1), Ivö, Sweden.
Blaksudden (Senon, Mammillatus
Zone), Ivö, Skåne, Sweden.
Blaksudden (Senon, Mammillatus
Zone), Ivö, Skåne, Sweden.
Blaksudden (site 1), Ivö, Sweden.
Identification
∑
Anomoeodus subclavatus
Anomoeodus subclavatus
Anomoeodus subclavatus
2
1
1
Indet. Pycnodontiformes
Alt: aff. Coelodus sp.
Anomoeodus sp.
1
Indet. Pycnodontiformes (not
described)
1
Anomoeodus subclavatus
1
Indet pycnodontiformes
(previously: Coelodus sp).
Anomoeodus subclavatus
11
Ischyodus sp.
1
Ischyodus sp.
2
Ischyodus sp.
1
cf. Enchodus sp.
1
Enchodus cf. E. gladidus
1
Mosasaur (Squamate)
Mosasaur (Squamate)
Mosasaur (Squamate)
Pliosaur
Pliosaur
Mosasaur (Squamate)
Ptychodus decurrens
1
1
1
1
1
2
1
Ptychodus mammilaris
1
Indet. Teleostei (not described)
Indet. Teleostei (not described)
Elasmobrachii (vertebrae)
1
1
1
Ischyodus sp.
1
Protosphyraena sp.
1
1
1
Appendix 1.2: Registered fossil specimens from RM PZ.
34
Identification
∑
Ivö Klack, Ivö, Sweden.
Anomoeodus subclavatus
1
27256
Ivö Klack, Ivö, Sweden.
Anomoeodus subclavatus
1
27257
27258
Ivö Klack, Ivö, Sweden.
Ullstorp, Kristianstad, Scania,
Sweden.
Ullstorp, Kristianstad, Scania,
Sweden.
Ullstorp, Kristianstad, Scania,
Sweden.
Ullstorp, Kristianstad, Scania,
Sweden.
Åsen (B. mammillatus Zone: Oyster
bank), Bromölla, Sweden.
Åsen (B. mammillatus Zone: Oyster
bank), Bromölla, Sweden.
Åsen (B. mammillatus Zone: Coquina
Bed and Oyster bank), Bromölla,
Sweden.
Åsen (B. mammillatus Zone: Coquina
Bed and Oyster bank), Bromölla,
Sweden.
Åsen (B. mammillatus Zone: Coquina
Bed and Oyster bank), Bromölla,
Sweden.
Ullstorp, Kristianstad, Scania,
Sweden.
Bäckaskog, Kristianstad, Scania,
Sweden.
Anomoeodus subclavatus
Protosphyraena sp.
1
1
Protosphyraena sp.
1
Indet. Pycnodontidae
2
Anomoeodus sp.
2
Indet. Teleost A.
6
Indet. Pycnodontidae
2
Category 1: Teleost vertebrae
3
Category 2: Teleost vertebrae
2
Category 3: Teleost vertebrae
4
Protosphyraena sp.
1
Indet. Ichthyodectiformes
1
PMU
Site and Biozone
27255
27259
27260
27261
27262
27263
27264
27265
27266
24768
27301
Appendix 1.3: Registered fossil specimens from the collection at PMU.
35
Appendix 2
Turtles
Aquatic Birds
Chondrichtyans
Actinopterygians
Faunal composition (%)
Dinosauria
Crocodilomorpha
Marine amniotes
0
10
20
30
40
50
Appendix 2.1: Bar chart showing the quantitative composition of vertebrates (stated in
percentage) present at the Kristianstad Basin (based on data compiled by Sørensen et al.,
2013; table 2, and this paper).
36
Appendix 3
Appendix 3.1: Unregistered specimen (crushing prearticular/vomerine tooth) of aff.
Coelodus sp. from Ullstorp. Orientation: A, occlusal view; B, basal occlusal view. Scale bar:
A-B = 0.4 mm (Collector & Photo: Tord Engfors – Ed: Mohamad Bazzi).
Appendix 3.2: Associated cycloid scales on white limestone matrix (elasmoid type with
numerous concentric circuli and radial rods) of indeterminable Ichthyodectiform fish from
Bäckaskog grustag (? B. mammillatus Zone), Scania, southern Sweden (PMU 27301: See
Fig. 4K). Descriptive remarks: The one figured specimen represents hitherto the best
preserved epidermal structure of bonyfish from southern Sweden. Geological remarks:
Bäckaskog grustag is currently not dated and the specimen lacks stratigraphic control.
Orientation: anterior view. Scale bar: 10 mm (Collector & Photo: Tord Engfors – Ed:
Mohamad Bazzi).
37
Appendix 4
Appendix 4.1: Saccular otolith (syn. Sagitta) of Teleostie incertae sedis from the Åsen
collection (LO. 11915t). Descriptive remarks: Large-sized otolith of uncertain systematic
affinity. Although is presumed to belong to a possible bottom-dwelling and/or mesopelagic
teleostean fish. Orientation: A, left inner face, B, anteriolateral views. Scale bar: A-B = 0.4
mm (Photo: Mohamad Bazzi).
38
Appendix 4.2: Unregistered teleostean fin spines from the Åsen collection. Scale bar: A-C =
0.4 mm (Collector: Tord Engfors - Photo: Mohamad Bazzi).
39
Appendix 4.3: Unregistered dental remains (incisiform and fang-like teeth) of uncertain
systematic affinity from Ullstorp. Descriptive remarks: Figured specimens B & D (Incisiform
teeth) might be belongings of the pycnodontiform fishes. Orientation: A-D, lingual view. Scale
bar: A-D = 0.4 mm (Collector: Tord Engfors - Photo: Mohamad Bazzi).
40