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On the Actinopterygian Fish Fauna (Upper Cretaceous: Campanian) from the Kristianstad Basin, Southern Sweden

On the Actinopterygian Fish Fauna (Upper Cretaceous: Campanian) from the Kristianstad Basin, Southern Sweden

Självständigt arbete Nr 88 On the Actinopterygian Fauna (Upper : ) from the Basin, Southern

Mohamad Bazzi On the Actinopterygian Fish Fauna (Upper Cretaceous: Campanian) Actinopterygian remains have been recovered from Upper from the , Cretaceous (uppermost lower to lowermost upper Campanian) marine strata of the Kristianstad Basin, southern Sweden. This Southern Sweden is the first record of Upper Cretaceous bony fish from the Fennoscandian shield. The fauna consists of higher taxa including (Pachycormidae), (Pachyrhizodontidae), (), (Enchodontidae), (Ichthyodectidae), and indeterminable fish. The ichthyofauna comprises a relative vast number of disarticulated meso-to macroscopic bony elements (e.g. Mohamad Bazzi vertebrae, isolated teeth, cycloid scales, , 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 assemblage of the Kristianstad Basin have resulted in major collection biases towards larger marine 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 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

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 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. Key words: Upper Cretaceous; Campanian; Osteichthyans; Actinopterygian; Teleostei; Palaeoecology; ; Kristianstad Basin; Åsen; Southern Sweden

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Table of content

1. Introduction ...... 1 2. Previous research...... 2 3. 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

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1. Introduction

Upper Cretaceous (Campanian) strata from the Kristianstad Basin, northeastern , Sweden, has yielded a plethora of 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 including a quantity of chondrichthyan (members, from both the Elasmobranch and clade) elements (e.g. mostly teeth). Also, osteichthyans (bony fish) and marine , such as plesiosaurs, , aquatic birds, dyrosaurid crocodilians, and unidentified taxa of cheloniid were major faunal components of this ancient marine (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, , bivalves (predominantly ) 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 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 ( subclavatus Agassiz, 1833), pachycormids ( sp. Leidy, 1857a), enchodontids ( gladidus Cope, 1872b), elopiformes (cf. 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 , but still they form a vital as well as a prominent portion of this Upper Cretaceous inner shelf- shallow marine environment.

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2. Previous research

Davis (1890) described and reported on several bony fish taxa from the and Early () of Scania, southern Sweden. Among the reported specimens, were pycnodontiformes (“” subclavatus, p. 415-416, pl. 42, fig. 16-18), (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. 1- 1a), 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 ( lundgreni Davis, 1890, p. 427- 428, 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 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 and stratigraphy (including maps) for the Kristianstad Basin. In brief summation, the Kristianstad Basin consists of a deeply weathered, crystalline ( 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 Sea, which during this interval covered a great portion of northwestern (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).

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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 bar shown in A-D is 1 km (after Bergström & Sundquist, 1978; Bergström, 1983a, b; Kornfält & Bergström, 1989).

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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 mammillatus (uppermost lower Campanian) and the Belemnellocamax balsvikensis (lowermost upper Campanian) belemnite (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 and/or lower Campanian affiliations (Friis & Skarby, 1981; Siverson, 1992a). The vertebrate fauna reported from Åsen comprises thirty-six , six bony fish, five ray, six and one 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 ( sp. personal observation) and 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 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 (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 , 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 fauna of Ivö has recently been extensively documented and comprises > 60 bivalvian, eight zooxanthellate and three azooxanthellate , seventeen gastropod, eighteen echinoid and sixteen asteroid, four belemnite and one ammonite, and twenty-five tube-dwelling 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 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 age (Christensen, 1998; Siverson & Cappeta, 2001). The onset of the Maastrichtian in northern Europe is marked by the appearance of 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 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 (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 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

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3.2), otolithes (Appendix 4.1), fin spines (Appendix 4.2), fragmented , 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 , 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 Åsen Ivö Klack Ignaberga Ullstorp Balsvik Scania Anomoeodus subclavatus x x x - - - Anomoeodus sp. - - - x x - Indet. Pycnodontidae x - - x - - Enchodus cf. E. gladiolus* - - - - - x cf. Enchodus sp. x - - - x - Indet. Ichthyodectidae x - - - - - cf. Pachyrhizodus sp. x - - - - - Protosphyraena sp. - x - x - - Indet. Teleost A. x - - - - -

*Scania = unknown locality

7. Systematic palaeontology

Class: Huxley, 1880 Cope, 1887 Regan, 1923 Order: Pycnodontiformes Berg, 1940 Family: Pycnodontidae Agassiz, 1833 sensu, Nursall 1996 : 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 ; 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.

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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 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; Poyato- Ariza & Wens, 2002; Kriwet, 2005a). As a morphological and ecological distinctive order of actinopterygians, the pycnodont 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 (Norian) to the (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 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 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 (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.

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Figure 4. Dental remains of stem-group (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.

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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 (), 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 (Arambourg, 1952), Netherlands (Goody, 1968), Belgium (Leriche, 1929; Friedman, 2012), England (Longbottom & Pattersson, 1987), (Kriwet et al., 2006), (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- 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 deposits () to the Palaeocene

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(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 (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 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, , 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 -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), (McNamarra et al., 1993) and Saudi Arabia (Kear et al., 2009).

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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 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).

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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 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 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 ), 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 (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 from the Upper Cretaceous of North America. They differ from the

18

North American 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 (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 (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 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

6

5 4 3 2

1 NUMBER OF TAXA TAXA OF NUMBER 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, latus, and 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 ) 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 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 85 % shark ∑

1000 15 % bony fish

800 378 259 600 169 17 400 55 Chondrichthyan teeth 31 0 Actinopterygian vertebrae 200 2 16 7 1 Total specimens Total Actinopterygian teeth 0 Coquina Bed Oysterbank Balsvikensis Balsvikensis green yellow

Actinopterygian teeth 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 . 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 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 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.

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40 34 35 30 25 20 Bulk sample 15 11 10 3 4 4 Numerical teeth count teeth Numerical 5 2 0 PYC ENC ICH PAC PAY Indet T Actinopterygians

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, 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.).

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 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 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 and finally I thank my good friend Filip Andersson for providing instructions in how to use Adobe Photoshop. 12. References

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13. Appendices

Appendix 1

LO Site and Biozone Identification ∑

LO. Åsen (Coquina Bed), Bromölla, cf. Pachyrhizodus sp. 2 11905t Sweden LO. Åsen (Balsvikensis yellow), Bromölla, cf. Enchodus sp. 1 11906t Sweden LO. Åsen (Balsvikensis green), Bromölla, cf. Pachyrhizodus sp. 1 11907t Sweden LO. Åsen (Balsvikensis green), Bromölla, Anomoeodus subclavatus 1 11908t Sweden LO. Åsen (Balsvikensis green), Bromölla, Indet. Ichthyodectidae 1 11909t Sweden LO. Åsen (Balsvikensis green), Bromölla, Indet. Pycnodontidae 3 11910t Sweden LO. Åsen (Balsvikensis green), Bromölla, Indet. Pycnodontidae 1 11911t Sweden LO. Åsen (Oyster bank), Bromölla, Indet. Teleost A. 5 11912t Sweden LO. Åsen (Oyster bank), Bromölla, Indet. Ichthyodectidae 1 11913t Sweden LO. Åsen (Oyster bank), Bromölla, cf. Pachyrhizodus sp. 1 11914t Sweden LO. Åsen (Coquina Bed), Bromölla, Teleostei incertae sedis 1 11915t Sweden (Saccular ) Appendix 1.1: Registered fossil specimens from the Åsen collection.

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RMPZ Site and Biozone Identification ∑

P.15753 Blaksudden (site 1), Ivö, Sweden. Anomoeodus subclavatus 2 P.13530 Blaksudden (site 1), Ivö, Sweden. Anomoeodus subclavatus 1 P.13746 Blaksudden (Senon, Mammillatus Anomoeodus subclavatus 1 Zone), Ivö, Skåne, Sweden. P.13745 Blaksudden (Senon, Mammillatus Indet. Pycnodontiformes 1 Zone), Ivö, Skåne, Sweden. Alt: aff. Coelodus sp. P.13670 Balsvik (Stora brottet), Anomoeodus sp. 1 Kristianstad, Skåne, Sweden. P.1416 Sweden (site 3) Sweden. Indet. Pycnodontiformes (not 1 (unknown locality) described)

P.13743 Blaksudden (Senon, Mammillatus Anomoeodus subclavatus 1 Zone), Ivö, Skåne, Sweden. P.13528 Sweden (Site 3) Sweden. Indet pycnodontiformes 11 (previously: Coelodus sp). P.13778 Ignaberga (site 1) Hässleholm, Anomoeodus subclavatus 1 Skåne, Sweden. P.13697 Blaksudden (Senon, Mammillatus Ischyodus sp. 1 Zone), Ivö, Skåne, Sweden. P.13770 Axeltorp (site 2) Bromölla, Skåne, Ischyodus sp. 2 Sweden. P.13698 Blaksudden (Senon, Mammillatus Ischyodus sp. 1 Zone), Ivö, Skåne, Sweden. P.13532 Balsvik (Norra brottet), cf. Enchodus sp. 1 Kristianstad, Skåne, Sweden. P.13685 Hedvigsdal (Stationen), Ystad, Enchodus cf. E. gladidus 1 Scania/Skåne, Sweden (locality is most likely wrong). R.1437 Blaksudden (site 1), Ivö, Sweden. Mosasaur (Squamate) 1 R.1438 Blaksudden (site 1), Ivö, Sweden. Mosasaur (Squamate) 1 R.1186 Blaksudden (site 1), Ivö, Sweden. Mosasaur (Squamate) 1 R.1298 Blaksudden (site 1), Ivö, Sweden. Pliosaur 1 R.1254 Blaksudden (site 1), Ivö, Sweden. Pliosaur 1 R.1125 Blaksudden (site 1), Ivö, Sweden. Mosasaur (Squamate) 2 P.1386 Annetorp (site 2), Malmö, Skåne, Ptychodus decurrens 1 Sweden (locality is wrong). P.1387 Annetorp (site 2), Malmö, Skåne, Ptychodus mammilaris 1 Sweden (locality is wrong). R.1237 Blaksudden (site 1), Ivö, Sweden. Indet. Teleostei (not described) 1 R.2217 Blaksudden (site 1), Ivö, Sweden. Indet. Teleostei (not described) 1 P.8697 Blaksudden (Senon, Mammillatus Elasmobrachii (vertebrae) 1 Zone), Ivö, Skåne, Sweden. P.13702 Blaksudden (Senon, Mammillatus Ischyodus sp. 1 Zone), Ivö, Skåne, Sweden. P.13583 Blaksudden (site 1), Ivö, Sweden. Protosphyraena sp. 1 Appendix 1.2: Registered fossil specimens from RM PZ.

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PMU Site and Biozone Identification ∑

27255 Ivö Klack, Ivö, Sweden. Anomoeodus subclavatus 1

27256 Ivö Klack, Ivö, Sweden. Anomoeodus subclavatus 1

27257 Ivö Klack, Ivö, Sweden. Anomoeodus subclavatus 1 27258 Ullstorp, Kristianstad, Scania, Protosphyraena sp. 1 Sweden. 27259 Ullstorp, Kristianstad, Scania, Protosphyraena sp. 1 Sweden. 27260 Ullstorp, Kristianstad, Scania, Indet. Pycnodontidae 2 Sweden. 27261 Ullstorp, Kristianstad, Scania, Anomoeodus sp. 2 Sweden. 27262 Åsen (B. mammillatus Zone: Oyster Indet. Teleost A. 6 bank), Bromölla, Sweden. 27263 Åsen (B. mammillatus Zone: Oyster Indet. Pycnodontidae 2 bank), Bromölla, Sweden. 27264 Åsen (B. mammillatus Zone: Coquina Category 1: Teleost vertebrae 3 Bed and Oyster bank), Bromölla, Sweden. 27265 Åsen (B. mammillatus Zone: Coquina Category 2: Teleost vertebrae 2 Bed and Oyster bank), Bromölla, Sweden. 27266 Åsen (B. mammillatus Zone: Coquina Category 3: Teleost vertebrae 4 Bed and Oyster bank), Bromölla, Sweden. 24768 Ullstorp, Kristianstad, Scania, Protosphyraena sp. 1 Sweden. 27301 Bäckaskog, Kristianstad, Scania, Indet. Ichthyodectiformes 1 Sweden. Appendix 1.3: Registered fossil specimens from the collection at PMU.

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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).

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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).

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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).

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Appendix 4.2: Unregistered teleostean fin spines from the Åsen collection. Scale bar: A-C = 0.4 mm (Collector: Tord Engfors - Photo: Mohamad Bazzi).

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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).

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