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Independent Project at the Department of Earth Sciences Självständigt arbete vid Institutionen för geovetenskaper

2015: 7

Disparity of Early Disparitet i Lamniformes Hajar från Tidig Krita

Fredrik Söderblom

DEPARTMENT OF

EARTH SCIENCES

INSTITUTIONEN FÖR

GEOVETENSKAPER

Independent Project at the Department of Earth Sciences Självständigt arbete vid Institutionen för geovetenskaper

2015: 7

Disparity of Lamniformes Sharks Disparitet i Lamniformes Hajar från Tidig Krita

Fredrik Söderblom

Copyright © Fredrik Söderblom and the Department of Earth Sciences, Uppsala University Published at Department of Earth Sciences, Uppsala University (www.geo.uu.se), Uppsala, 2015 Sammanfattning

Disparitet i Lamniformes Hajar från Tidig Krita Fredrik Söderblom

Morfologisk disparitet är ett mått på hur stor utsträckningen av morfologisk variation är. Detta mått räknas ut genom att jämföra landmärken utplacerade på bilder av föremål som ska undersökas. I detta projekt undersöktes den morfologiska dispariteten hos tänder från håbrandsartade hajar (Lamniformes) under tidig krita. Att just deras tänder undersöktes beror på att den större delen av hajars skelett är gjort av brosk vilket lätt bryts ned efter djuret avlidit. Deras tänder är dock gjorda av ben vilket har lättare att bli bevarat som . Utöver detta så kan formen på tänder beskriva djurs födoval och levnadssätt. Gruppens tänder undersöktes därför även för att belysa eventuella förändringar i diet och ekologi under tidig krita. Resultatet av denna analys visar på en expansion av tandform under denna period från långa och smala tänder under Barremium till en större variation under Albium där även mer triangelformade och robusta tänder dyker upp. Detta har tolkats som en adaptiv artbildningsperiod för gruppen då både nya byten (t.ex. teleostfiskar och havs- sköldpaddor) diversifierade och uppkom samtidigt som vissa marina predatorer (ichthyosaurer och plesiosaurer) minskade i antal under denna tidsperiod. Detta ändrade troligen de selektiva trycken på håbrandsartade hajars tandmorfologi samt lämnade ekologiska nischer öppna som dessa kunde anpassa sig till vilket i sin tur ledde till expansioner i morfologisk disparitet, diet och ekologi.

Nyckelord: Lamniformes, disparitet, tidig krita, morfologi, morfometri

Självständigt arbete i geovetenskap, 1GV029, 15 hp, 2015 Handledare: Nicolàs Campione Biträdande handledare: Benjamin Kear Institutionen för geovetenskaper, Uppsala universitet, Villavägen 16, 752 36 Uppsala (www.geo.uu.se)

Hela publikationen finns tillgänglig på www.diva-portal.org

Abstract

Disparity of Early Cretaceous Lamniformes sharks Fredrik Söderblom

The geological range of lamniform sharks stretches from present day such as (great white ) back to the at the moment oldest undoubted fossil finds during the Early Cretaceous. In this paper a geometric morphometric analysis was performed on images of Early Cretaceous lamniform teeth collected from published literature in to examine the change in disparity (range of morphological variation within a group) throughout the time period. Due to limited availability of published material and time constraints only the and ages were investigated. The Barremian exhibited tall and narrow morphologies while the Albian showed a wide range of morphological variation including more robust, wide and sometimes triangular shapes but also displayed further specialization of the tall and narrow forms. This change is likely indicative of a dietary and ecological expansion from only eating for example small and soft- bodied creatures to a wide range of prey for the group, including larger and more robust such as marine and large bony fish. This in combination with the decline of some marine predators as well as the diversification of possible prey is interpreted as that an adaptive radiation of the Lamniformes could have taken place during the latter half of the Early Cretaceous.

Key words: Lamniformes, disparity, Early Cretaceous, morphology, morphometric

Independent Project in Earth Science, 1GV029, 15 credits, 2015 Supervisor: Nicolàs Campione Co-Supervisor: Benjamin Kear Department of Earth Sciences, Uppsala University, Villavägen 16, SE-752 36 Uppsala (www.geo.uu.se)

The whole document is available at www.diva-portal.org

Table of contents 1. Introduction 1 2. Materials and methods 2 3. Results 3 4. Discussion 5 4.1 Discussion of the results 5 4.2 Dental adaptations and habitats 6 4.2.1 The Barremian 6 4.2.2 The Albian 8 4.3 On changes in disparity, diversity, diet and feeding strategies 10 5. Conclusions 13 6. Acknowledgements 14 7. References 14 7.1 Printed refereces 14 7.2 Internet references 17 8. Appendix 17

1. Introduction

Chondrichthyes is the name of a group including the clades () as well as (sharks and rays) (Miller et al., 2003; Hickman et al., 2011). Fossil scales prove chondrichthyans to possibly have existed since the Late (Janvier, P., 1996 & Turner, S. in Miller et al., 2003). They are fish with a skeleton made of , a feature that developed as they evolved out of their ancestors with skeletons made of (Hickman et al., 2011). The fact that their teeth are better mineralized than the rest of their skeleton is the reason that teeth are one of the most common kind of fossil found belonging to them since cartilage is less likely to survive the fossilization process (Shimada, 2005, 2007; Whitenack & Gottfried, 2010). Sharks that are a part of them have been determined to have existed during the Early on the basis of fossil teeth (Miller et al., 2003) and even back to the Lower Siluran on the basis of simple placoid scales (Karatajūté- Talimaa, V., 1973, 1992 in Sansom et al., 1996). The clade Lamniformes is an order of sharks that is present on Earth today and includes the great (Carcharodon carcharias) (Cappetta, 2012). The order’s origin was however during the Early Cretaceous with one of the earliest undoubted fossil finds belonging to the early part of the Late (Rees, 2005). Although the possibility of an origin during the has also been proposed (Underwood, 2006). The Cretaceous is a time period that spans from 145-66 million ago (Ma) (Cohen et al., 2015). The period is divided into the Early Cretaceous (145- 100.5 Ma) and the (100.5-66 Ma) (Cohen et al., 2015). The Early Cretaceous was the focus of this study. It can be further subdivided into the (145-139.8 Ma), the Valanginian (139.8-132.9 Ma), the (132.9- 129.4 Ma), the Barremian (129.4-125 Ma), the (125-113 Ma) and the Albian (113-100,5 Ma) (Cohen et al., 2015). During the beginning of the Early Cretaceous the global climate became more arid than before (Benson & Druckenmiller, 2014) (evidenced in part by clays having an increased proportion of the mineral smectite (Weissert & Channell, 1989; Hallam et al., 1991)), the ocean surface became more oligotrophic (Danelian & Johnson, 2001; Tremolada et al., 2006 in Benson & Druckenmiller, 2014) and temperatures rose to eventually reach a high approximately 100 million years ago (Gould et al., 2001). Radiations of several different organisms took place in the Cretaceous and many of them originated during the Early Cretaceous (Sadava et al., 2011). The first angiosperms (flowering plants) appeared during the Early Cretaceous (Gould et al., 2001; Sadava et al., 2011), triggering a radiation of insects such as butterflies, bees, moths, as well as ants (Gould et al., 2001). The first snakes arose during the Early Cretaceous (Benton, 2005; Sadava et al., 2011). Other organisms such as gymnosperms, , sharks, plesiosaurs, belemnites, ammonites (Gould et al., 2001), (Gould et al., 2001; Sadava et al., 2011), (Gould et al., 2001) (pterodactyloid pterosaurs during the Early Cretaceous (Benson & Druckenmiller, 2014)) and turtles (Early Cretaceous) (Scheyer et al., 2014) would diversify during the Cretaceous. Large predators such as also appeared in the Cretaceous as well as the first true and (Gould et al., 2001). With this paper a dataset connected to images of Early Cretaceous Lamniformes shark teeth will be presented (see appendix). A geometric morphometric analysis of the teeth in these images was also performed. The results of this analysis will be presented as diagrams. A morphospace diagram showcasing

1 the measure of morphological variation between specimen in a mathematical space and a disparity through time diagram showing the disparity (the range of morphological variation within a group (Briggs & Crowther, 2001)) of lamniform teeth during the different ages of the Early Cretaceous. A t-test was also performed to account for the significance of the sampled specimen in question between the different time bins. The hypothesis being tested is that since a sizeable amount of radiations seem to have taken place at the time investigated in this paper, the results of the analysis might show a high amount disparity and possibly an increase in morphological variation during the aforementioned time period.

2. Materials and methods

A total of 146 images of Lamniformes teeth specimen were located using published literature available in both digital and physical formats and compiled into a dataset (see table 1 in appendix). The preferred view in the images was labial (seen from the outside of the mouth), however if this was not available then lingual view (seen from the inside of the mouth) was used. The selection of the images gathered and used was based on availability and unfortunately somewhat restricted by amount of time available to work on this study. These images were then scanned (if the image was from a physical source) or saved (if the source was digital) as JPEGs. At the same time information about the images such as where it was taken from, which page it was taken from, figure number, which dental unit it was from, what specimen number it had, what order, , and species it was, what view the images was taken in, its relative position in the cranium of the individual, what continent, country, locality and formation it was found in as well as what era, period, and age it belonged to. This information was put together into a dataset in Microsoft excel. The images were then cropped using Adobe Photoshop CC 2014 (Adobe Systems Software Ireland Ltd) and a scale bar was also moved into the newly created image if it was available in the original figure. The teeth were then re- oriented by flipping them in Microsoft Paint so the apex of the crown (see figure 7 in appendix for explanation of tooth component locations) in every image was oriented either straight up if the tooth crown was straight or up and to the left if the tooth crown was inclined. A tps file was created by inputting the folder containing only the images of complete specimens (85 in total (see table 2 in appendix) of which seven were of Barremian age and 78 were of Albian age) in the open-source program tpsUtil (version 1.60) created by F. James Rohlf (Rohlf, 2015). Afterward in the program tpsDIG2 (version 2.18) created by F. James Rohlf (Rohlf, 2015) the images of the tps file were placed on a Euclidian coordinate grid. tpsDIG2 was then used to digitize the images by placing semilandmarks along the edge of the crown going from the left side of the crown to the right (see figure 7 in appendix). The semilandmarks made up a curve and were then all resampled to 74 points along the curve with equal length between each other. In notepad, the semilandmarks in the tps file were changed into homologous landmarks. This file was then entered into tpsUtil where it was used to create a sliders file in which all landmarks were linked and marked to slide except for the first and last ones. The tps- and sliders files as well as the dataset (table 2 in appendix) were then imported into the statistical programming program RStudio (RStudio) using the package geomorph (a package that can analyze the shape of

2 both two-dimensional and three-dimensional landmark data) (Adams, D. C. & Otarola-Castillo, E., 2013; Adams et al., 2015) where they then were used to run a generalized procrustes analysis (GPA) (using the tps- and sliders files all the specimens were moved to the same point in mathematical space, scaled to approximately the same size and rotated in an as close as possible same direction so that distances between the same landmark in different specimen could be measured) (Adams et al., 2015)) and a principal component analysis (PCA) (plots the specimens aligned in the GPA along principal axes (Adams et al., 2015)) that generated a morphospace diagram of the sample (figure 1). The samples (dots) were then partitioned into time bins corresponding to their geological age using the imported dataset and the RStudio which () command. The dots on the edges of the corresponding time bins were then linked by lines using a function called minimum convex hulls (Campione, N.) and were colored by using the RStudio command points (). Images of the mean of principal component 1 (PC1) and principal component 2 (PC2) minimum and maximum values were generated in RStudio. These were then added to their respective positions in the morphospace diagram by using the program Inkscape (Inkscape). The results from the PCA were then used with the disparity () command from the RStudio package geiger (Harmon et al., 2008) to calculate the disparity for the time bins in question and then a diagram of the disparity through time was plotted. A t-test was also performed on the principal component scores obtained from the PCA in RStudio to test for statistical significance in the sample that was analyzed and the mean of the two ages.

3. Results

The results of the analyses are presented below and include the morphospace diagram (figure 1), disparity through time diagram (figure 2) and the results of the t- test. The morphospace in figure 1 does not have any units since variables such as size and angle were removed during the analysis. The differences between landmarks were measured using the Euclidian coordinate grid the images were placed upon. Principal component 1 (PC1) (figure 1) represents the greatest morphological difference of one and the same landmark between all specimen in the analysis. Principal component 2 (PC2) (figure 1) represents the second to greatest morphological difference in another landmark between all specimen used in the analysis.

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Figure 1. Morphospace diagram showing the extent of morphological variation during the ages included in this sample. Samples of Barremian age are shown as red and samples of Albian age are shown as blue. The images on the sides of the diagram show minimum and maximum extremes of the samples’ shape along that particular principal component axis. Principal component axis 1 shows the height and width change in morphology inside the morphospace diagram from narrow and tall (left side) to low and wide (right side). Principal component axis 2 shows the degree to which the cusplets are present where the top part of the figure has large cusplets and the bottom of the figure has none. Furthermore the width of the base of the crown increases in size from the top left toward the bottom right of the morphospace diagram.

Figure 2. Diagram showing the disparity (range of morphological variation within a group) (y axis) through time (x axis) of the sampled specimens. Specimens from other stages were not included in the analysis and are therefore not shown in this diagram.

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In the T-test between the mean of the Barremian specimen (calculated to be - 0.14433504 on the principal component 1 axis (figure 1)) and the mean of the Albian specimen (calculated to be 0.01295314 on the principal component 1 axis (figure 1)) a p-value = 0.0307 was obtained.

4. Discussion

4.1 Discussion of the results

The results of the morphospace diagram (figure 1) show an increase in disparity as does the disparity through time diagram (figure 2). However three things are worth taking note of. The first problem with these results is that the Aptian, the age which should be placed in between the Barremian and the Albian is missing. This leads to a problem since the statistical significance of the samples reflecting reality cannot be properly ascertained in the detail that would be desirable. In other words, an increase of disparity can be seen in both figure 1 and figure 2 but since there is no accounting for the disparity during the time that is missing (the Aptian), there is no knowing for sure whether there was an increase, decrease or no change in disparity of lamniform teeth from the Barremian to the Aptian or from the Aptian to the Albian. Although what can be seen in both figure 1 and 2 is that an overall increase in disparity of lamniform teeth from the Barremian to the Albian seems to have taken place. So at least in the Albian lamniforms seems to have had a wider range of morphological variation than in the Barremian. The second thing that is noteworthy is that since the Barremian only had seven specimens attributed to it in the analysis it is possible (and also quite likely) that this does not closely enough represent the true extent of Barremian lamniform tooth morphology (and would therefore not show if the true disparity was initially high or not as was hypothesized). There is also the possibility of a sampling bias considering the small amount of Barremian specimen and this would affect the interpretation of the results where it could look like there was a greater expansion in the range of morphology than there actually was during the time from the Barremian to the Albian. All the Barremian samples used in the analysis were of the same family (Eoptolamnidae) and consisted of only three species and genera (Eoptolamna eccentrolopha, stychi sp. nov. and Protolamna sarstedtensis sp. nov.) (table 2 in appendix). In the complete dataset (table 1 in appendix) 22 specimens are of Barremian age. 19 of these belong to the family Eoptolamnidae. The three remaining Barremian specimen were not assigned to a particular family in the published literature they were taken from. However tooth morphology can vary on the level of genera/species to different degrees and even vary in a single individual’s mouth (heterodonty) depending on not only its belonging in a certain genera or species but also changes such as those during ontogeny (changes taking place when an individual grows up) and (gender differentiated traits) (Cappetta, 2012). The diet of a shark can also be influenced by the tooth morphology it possesses since this determines what prey it will be able to catch and consume (Cappetta, 2012). Other than the Eoptolamnidae families like Cretoxyrhinidae and Odontaspidae were also present during the Barremian (Underwood, 2006). Although this is the case, it is possible that these families either could have possessed a similar dental morphology, adapted to a similar diet or a completely different morphology adapted for other kinds of prey. Measured disparity is more affected by

5 the addition of new specimen or removal of existing specimen at the edges of that time bin’s area since this would change the appearance of the time bin in morphospace to a greater degree than adding or removing a specimen in the middle of the time bin’s area (Foote, 1993). Also, the larger the size of a group within a time bin in morphospace the more it affects disparity (Foote, 1993). In other words, if a group is large and very peripheral in morphospace it affects that time bin to a much greater extent than a small and centralized group within time bin.This would mean that if Barremian specimen with quite different morphologies than the ones in the morphospace diagram were added this would likely expand the Barremian time bin’s area. However if the new specimen added were very similar morphologically to the ones already in the time bin this would likely not expand the time bin or increase disparity to any larger degree. In other words, this does not completely invalidate the Barremian time bin shown in figure 1 although it should be expanded upon with more specimens to more correctly represent the true extent of lamniform tooth morphology during this particular age. The Albian seems to be quite well sampled and is showing a quite varied morphology as seen in both figures 1 and 2. It should be noted though that 25 of the 78 specimens from the Albin time bin in the analysis were defined by the authors of the paper they were taken from as belonging to either the latest Late Albian or earliest Early (Siverson et al., 2013) and therefore should be considered as a possible source of error in the analysis performed in this paper since they might belong to an age not investigated here. Both the Barremian and Albin time bins will be discussed below. Lastly, the results of the t-test showed a p-value of 0.0307 which means in turn that the sample is significant at the 5% level (Olsson et al., 2005). Furthermore the t-test also showed the mean of the Barremian specimens to be located at - 0.14433504 on PC1 and the mean of the Albian at 0.01295314 on PC1. If compared to figure 1 this seems to be correctly assessed and would likely serve as evidence for a shift in average morphology of the two time bins. The Barremian time bin’s PC1 average on the negative (left) side of the PC1 axis suggests a tall and narrow morphology, most likely more adapted to catching and consuming soft-bodied animals or smaller fish. The Albian time bin’s PC1 average is placed in a more central location along the PC1 axis, which would indicate a more varied (and wide) range of morphologies (as also evidenced by the negative and positive extremes of specimen in the Albian time bin) including the narrow and tall morphology already present in the Barremian but also more low and wide teeth useful for processing more robust prey. Descriptions of dentition types as well as their implications concerning diet, examples of feeding strategy and some possible driving forces behind the dentition morphology shifts of lamniforms during the Early Cretaceous will be discussed below (subheadings 4.2 and 4.3).

4.2 Dental adaptations and habitats

4.2.1 The Barremian

The disparity of the Barremian specimens shown in figure 1 seem to indicate a tooth morphology with a quite straight (except for one of the specimen which seems to have its main cusp bent slightly more than the others toward the rear of its mouth), tall, narrow and pointed main cusp and with somewhat medium sized cusplets (smaller thorn-like structures next to the main cusp (see figure 7 in appendix)). The

6 teeth seem to be either of the clutching or tearing type dentitions mentioned by Cappetta (2012). Clutching type dentition (figure 3) generally involves quite small teeth set in rows with a smooth/straight or either lingually or labially bent crown (Cappetta, 2012). Clutching dentition often includes one or more pairs of cusplets and is usually an adaptation that is used by present day elasmobranchs to puncture and hold on to (aided by the cusplets) softer prey such as fish and that move very quickly in open (Cappetta, 2012). Ramsay & Wilga (2007) did however show that the recent shark Chiloscyllium plagiosum (white-spotted bamboo shark) is physically capable durophagy (eating hard-shelled prey, e.g. and hard-shelled mollusks) by rotating its teeth lingually (toward the inside of the mouth) through the use of dental ligaments and by doing so changing the function of its clutching dentition into that of a crushing dentition. If this function existed in Early Cretaceous lamniform sharks though is unknown. Furthermore, Cappetta (2012) also mentions that this type of dentition is common in sharks benthic and demersal zones (living on respectively a bit above the bottom of the sea) and also in the neritic zone (above the continental shelf).

Figure 3. Simplified example of the range of clutching type dentition present in this paper.

Tearing type dentition (figure 4) usually has one to several pairs of lateral cusplets (though in some genera they are missing) and more noticeable cutting edges along the side of the crown (Cappetta, 2012). Some sharks with tearing type dentition even acquired a cutting function during the Early Cretaceous due to increased sharpness of the crown’s edges (Cappetta, 2012). Sharks with this dentition possess the ability to bite through bone (Ramsay & Wilga, 2007) and are therefore more likely to eat more robust prey than the clutching dentition mentioned above. The habitats of extant sharks with tearing type dentition include bathyal (deep ), littoral (near shore), epipelagic (far up in the open ocean outside the continental shelf) and pelagic (open ocean) environments (Cappetta, 2012).

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Figure 4. Simplified example of the range of tearing type dentition present in this paper. Tearing type dentition may also include more noticeable cusplets.

4.2.2 The Albian

Figure 1 seems to indicate that during the Albian stage, morphology was quite varied within the Lamniformes order. The Albian time bin seems to have expanded a lot in the positive direction and only a little in the negative direction on PC1. It also expanded a lot in the negative direction but only somewhat in the positive direction on PC2. The order seems to have increased its dental disparity in the Albian compared to the Barremian by (except for the morphologies present then) either (1) shortening the main cusp (and either keeping it straight or bending it), widening the crown base and elongating the cusplets (specializing more in clutching dentition), (2) elongating its main cusp as well as decreasing the size of its cusplets (and in some cases widening the crown base) or (3) bending the apex of its main cusp toward the rear to varying degrees, widening the crown base and losing its cusplets (figure 7 in appendix and figure 1). This wide a range of morphological expansion in approximately 12-25.5 million years would indicate some sort of adaptive radiation due to a change in selective pressures (more on this below in subheading 4.3). Again following the terminology used by Cappetta (2012), the tearing- and clutching type dentitions mentioned previously (see 4.2.1 The Barremian) also seem to be present in this Albian (see figure 1) and seem to have somewhat further specialized. Additionally, types such as sensu stricto cutting dentition and cutting-clutching dentition are also encompassed by the Albian time bin in figure 1. Both of these are subtypes of the cutting type dentition (Cappetta, 2012). Sensu stricto cutting dentition (figure 5) is characterized by labio-lingual flattening and also a general widening (Cappetta, 2012). Because of this labio-lingual flattening and the fact that this dentition type’s crown often is bent backwards (though sometimes it is straight) and linked together (complete linkage is not always present) into a continuous row of teeth creates something akin to a blade (Cappetta, 2012). Teeth with this kind of dentition may also have serrated edges (Cappetta, 2012). Sensu stricto cutting dentition is able to cut the flesh of prey into pieces as well as chew through bone by acting as a pair of scissors (Ramsay & Wilga, 2007) allowing it

8 to eat very robust and large prey. Sharks with sensu stricto cutting dentition can often be found in bathyal, pelagic and neritic areas of the sea (Cappetta, 2012).

Figure 5. Simplified example of the range of sensu stricto cutting type dentition present in this paper.

The cutting-clutching subtype dentition (figure 6) exhibits a quite strong differentiation between the upper and lower jaw of the individual (Cappetta, 2012). One jaw has high and narrow cusps (most often in (but not restricted to) the anterolateral and anterior teeth) (Cappetta, 2012). The other jaw can have wide teeth that are flattened labio-lingually (Cappetta, 2012). Like sensu stricto cutting dentition, cutting-clutching dentition can also be used to cut through bone and flesh of more robust creatures. Cutting-clutching dentition can be found in sharks living in bathyal, epipelagic and littoral environments (Cappetta, 2012). It should be noted that Ramsay & Wilga (2007) do mention in their paper that cutting dentition seems to be present in sharks that nourish themselves by consuming fish, mammals and soft- bodied invertebrates.

Figure 6. Simplified example of the range of cutting-clutching type dentition present in this paper. Left side shows the range of cutting teeth. Right side shows the range of clutching teeth (most often present in the anterior and anterolateral teeth of one jaw).

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4.3 On changes in disparity, diversity, diet and feeding strategies

The hypothesis posed in the introduction of this study would seem to fit quite well with the results of the analyses that were conducted. However since as mentioned earlier only the latter part of the Early Cretaceous was covered by this study, the Barremian stage was represented only by seven specimen and there was a hiatus of specimen during the Aptian in this study the correctness by which the results actually depict the change in disparity from the Barremian to the Albian is not quite optimal. Expanding upon the dataset and running additional analyses would be advisable and would have been done if not due to time constraints. Nevertheless the Albian time bin had a decent amount of specimens attributed to it and the results during that particular stage should be viewed as a quite correct representation of the disparity calculated for that interval of time. If one would assume that the Barremian time bin is somewhat representative of the actual disparity of that stage and then compared it to the Albian then the increase in morphological variation would certainly support the sizeable radiations mentioned in the introduction of this paper. Since the Lamniformes seemed to expand into cutting dentition (except for the tearing and clutching dentitions present in both the Barremian and the Albian) capable of through bone and flesh of more robust animals than small fish or for that matter soft invertebrates there would likely have been some event(s) that triggered this push toward a dentition of wider and more sturdy blade-like teeth. By the Early Cretaceous teleost fishes had already appeared long ago however they diversified during this time (Gould et al., 2001; Benton, 2005). The elopomorphs (clade including eels and their relatives) and clupeomorphs (clade including extant anchovy and ) arose in the Early Cretaceous (Benton, 2005). It is worthy to mention that during the Early Cretaceous the (superficially -like marine reptiles) that preyed on fish and cephalopods (Gould et al., 2001) throughout the majority of the had started to decline (Gould et al., 2001; Maxwell & Kear, 2010) and would eventually go extinct a time after the end of the Early Cretaceous (the latest finds are from the Cenomanian, the first age of the Late Cretaceous) (Maxwell & Kear, 2010). It was toward the end of the Early Cretaceous that the up to 4.2 meter long predatory fish Xiphanctinus (Benton, 2005) made its appearance (Gould et al., 2001). fossil have been found with 1.6 meter long prey in its stomach (Benton, 2005). This increase in fish taxa (and in some cases size) might have been brought upon by the decline and of the ichthyosaurs. Their disappearance might have left an open role in Cretaceous ecosystems that would be filled by Xiphactinus and other fish (possibly including some sharks). Although Xiphactinus might seem large, some species of shark from a family called the Cretoxyrhinidae (one of the lamniform families included in the analysis) could reach a size of six meters long, weigh an approximate of 1.5 tons (Gould et al., 2001) and were equipped with either tearing (Benton, 2005) or cutting dentition (Shimada & Hooks, 2004; Siverson et al., 2013). That the cretoxyrhinid sharks have preyed upon Xiphactinus is supported by the work on Late Cretaceous mantelli from the Niobrara Chalk () done by Shimada (1997) where a well preserved fossilized skeleton of C. mantelli was found with the remains of a Xiphactinus audax located posterior to its head. Even plesiosaurs are suggested to possibly have been eaten by Cretoxyrhina from the discovery of what could be

10 plesiosaur gastroliths (stones ingested by animals to help break down food often only found in marine low energy sediments in association to plesiosaur ) along with Cretoxyrhina remains (Shimada, 1997). However no plesiosaur remains were found associated with the gastroliths. Shimada (1997) also mentions attacks on mosasaurs by cretoxyrhinid sharks where one of the mosasaurs show evidence of an infection and subsequent healing around an embedded cretoxyrhinid tooth. This was interpreted as evidence of the cretoxyrhinid shark employing an active feeding strategy (as opposed to scavenging a carcass). Siverson (1992 in Schwimmer et al., 1997) hypothesized that C. mantelli could have fed on sea turtles but the only evidence for this was body size and dental morphology. This idea was later confirmed by Shimada & Hooks (2004) through what was determined to be C. mantelli teeth embedded into an individual of the Late Cretaceous protostegid species gigas, a species with a carapace (upper part of a turtle’s shell) that could reach lengths of more than two meters. The were a turtle family with a geological range from the Albian (last age of the Early Cretaceous) to the Late (Late Cretaceous) (Shimada & Hooks, 2004). They were part of the turtle superfamily Chelonioidea that appeared as turtles diversified during the late Early Cretaceous (Scheyer et al., 2014) and unlike many of their more ancient relatives the chelonioids were more adapted to open marine environments (Scheyer et al., 2014). The appearance of these new open marine turtles could have changed the selective pressures on lamniform tooth morphology from the more narrow clutching type (often found in neritic or benthic conditions suitable for catching smaller fish and soft-bodied prey) or tearing type dentitions present in the Barremian time bin toward the more robust and wide tearing type or cutting type (found in both pelagic, neritic and bathyal areas) dentitions appearing in the Albian time bin. While C. mantelli was of a species and time not investigated in this paper, there is the possibility that the cretoxyrhinids (or for that matter other lamniforms) investigated in this paper could have inhabited a similar ecological role as (or at least high trophic level predator) of some Early Cretaceous ecosystems. This might be somewhat supported by Schwimmer et al. (1997) stating that recent and fossil shark dentition belonging to sharks of the superorder Galeomorpha (including the order Lamniformes) has been very similar since the Early Cretaceous and therefore a similar feeding behavior can be assumed. Except for this Schwimmer et al. (1997) also mention that predatory behavior has always been common in sharks of large size. Teeth belonging to one of the cretoxyrhinids used in the analysis of this paper were of the species Cretoxyrhina vraconensis. This species has been estimated to have reached a length four meters as an adult (Siverson et al., 2013). As a juvenile its anterior teeth were narrow and well adapted for catching cephalopods, bony fish and small elasmobranchs (Siverson et al., 2013). Through ontogenic changes the adult individuals would instead have a more triangular and wide cutting dentition, feeding upon larger prey (Siverson et al., 2013). The size and dentition of C. vraconensis would seem to indicate an ecological role somewhat similar to that of C. mantelli. However, the C. vraconensis teeth used in the analysis were described by Siverson et al. (2013) as being from a top of the food chain predator in a coastal marine environment where large marine reptiles have not been found. This would not preclude the possibility of C. vraconensis living in open marine environments though (however this is just speculation). As mentioned earlier, both sharks with tearing type dentition as well as cutting type dentition can be found in open marine (pelagic) settings (Cappetta, 2012).

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The genus is also one of the genera included in the analysis performed in this paper. This genus ranges from the Albian (latest age of the Early Cretaceous) to the (latest age of the Late Cretaceous) and possesses cutting type dentition (Schwimmer et al., 1997). It has been suggested and discussed at length by Schwimmer et al. (1997) that members of the genus used this cutting dentition primarily for scavenging on carcasses from animals such as mosasaurs, plesiosaurs, marine turtles and teleost fishes (as well as occasionally some terrestrial that had died and were transported by water out into the ocean). Many of these organisms are believed to have been larger than Squalicorax. Therefore it is believed that they were likely deceased when being fed upon. Schwimmer et al. (1997) mentioned that this might not necessarily indicate a feeding strategy of obligate scavenging but seemed to indicate that it was most likely the primary means used to obtain sustenance by the members of the genus. This use of cutting dentition by Squalicorax shows that just because the dentition could be used to actively hunt and consume large/robust prey (as for example it was likely used by members of the Cretoxyrhinidae), it doesn’t mean that active hunting necessarily was the means used by every wielder of the dentition to obtain food. Another example of scavenging (or possibly actively attacking prey) is described by Shimada et al. (2010) where teeth (most similarly described as the cutting-clutching dentition in this paper) from six to seven individuals (1.5 to 4.2 meters long) of Late Cretaceous (also known as Cretolamna) appendiculata (an Albian specimen of Cretolamna sp. was used in the analysis of this paper (table 2 in appendix) and seems to have similar dentition to the teeth of Cretalamna appendiculata described by Shimada et al. (2010) which means a similar diet could be inferred) were found both next to and imbedded into a 6.4 to 9.2 meter long individual of the plesiosaur species Futabasaurus suzukii. Plesiosaurs have been shown to have declined in morphological disparity from a high in the middle of the Jurassic (pliosauroid plesiosaur decline in the ) to a low in the early part of the Early Cretaceous (Berriasian-Barremian) (seemlingly substantial cryptoclidid plesiosauroid disparity loss around the Jurassic-Cretaceous boundary) (Benson & Druckenmiller, 2014). Part of them thereafter increased their range of morphological variation (, Leptocleididae and of the Plesiosauroidea), elevating plesiosaur disparity once again toward a high level in the late Early Cretaceous (Aptian-Albian) (Benson & Druckenmiller, 2014) suggesting that they too radiated during the (late) Early Cretaceous. The decline and subsequent radiation of these animals could have acted as another trigger for the radiation and increase in tooth morphology diversity of Lamniformes (as well as possibly bodysize?) by leaving open ecological roles available to adapt into and thereafter giving lamniform sharks additional prey to feed on in the form of smaller sized species of plesiosaurs or young/old, diseased and deceased individuals of large plesiosaur species. The fossil remains of the plesiosauroid described by Shimada et al. (2010) showed no evidence of healing though so it is believed that the biting likely took place after the death of the . Another idea mentioned by Shimada et al. (2010) is that the sharks attacked the plesiosaur with a fatal outcome. However the definite cause of the plesiosaur’s death was not established. Lastly the diversity of lamniform genera (which can possibly but does not necessarily have to be related to changes in morphological variation and dentition types) can also be mentioned through which genera were present during the time analyzed in this paper (Barremian-Albian). During the Barremian at least four lamniform genera seem to have been present (Hispidaspis, Protolamna (Cappetta,

12

2012), Leptostyrax (Schmitz et al., 2010), Eoptolamna (Kriwet et al., 2008)). The Aptian contains at least eight genera (, , Hispidaspis, Paraisurus, Protolamna, Eostriatolamna (Cappetta, 2012), Leptostyrax (Schmitz et al., 2010), (Shimada, 2007)). The Albian seems to be where lamniform genera of the late Early Cretaceous were the most abundant in numbers with 20 genera present (Anomotodon, Scarpanorhynchus, Hispidaspis, Johnlongia, , Pseudoisurus, Acrolamna, , Cretoxyrhina, Paraisurus, Leptostyrax, Protolamna, Pseudoscarpanorhyncus, Squalicorax, , Dwardius, Eostriatolamna (Cappetta, 2012), Priscusurus (Kriwet, 2006; Cappetta, 2012), Carcharias (Siverson, 1997; Everhart, 2009), Cretolamna (Siverson, 1997; Cappetta, 2012)). There seems to be an increase in genera from the Barremian to the Aptian and then also an increase from the Aptian to the Albian. Although it is possible that these increases could also be driven by the lack of study on shark fossils from pre- Albian Cretaceous rocks (though especially earliest Cretaceous rocks) (Underwood, 2006) and the Barremian and Aptian therefore not having been represented fully. If it would be assumed however that the Barremian and Aptian had been examined well enough, this increase in genera by itself does not mean that the range of morphological variation (disparity) increased. This would only tell the change in number of genera through time and does not tell if there was an overall varied or very similar morphology present throughout the group during the Barremian to the Albian (and therefore does not say anything about the ecology of these sharks either). In other words, high diversity in genera does not have to mean high disparity. When compared to the morphospace diagram in figure 1 and disparity through time diagram in figure 2 though the diversity and the disparity both seem to indicate a time of adaptive radiation for the Lamniformes order that took place sometime during the (late?) Early Cretaceous (possibly in the Aptian and/or Albian).

5. Conclusions

Following the pattern of many other organisms during the Early Cretaceous the Lamniformes order of sharks likely radiated in an adaptive manner as well. Although the range of their morphological variation early on during this time could not be confidently assessed, their disparity did increase toward the end of the Early Cretaceous, with a radiation possibly having taken place during the Aptian and/or Albian. The combined action of large marine predator decline (such as ichthyosaurs and plesiosaurs), subsequent open ecological niches and radiation of prey (e.g. open marine turtles, teleost fishes) likely spurred on selective pressures which changed the tooth morphology of lamniforms. The tall and narrow clutching- or tearing type dentitions during the Barremian shifted toward a very varied morphology including clutching-, tearing-, sensu stricto cutting- and cutting-clutching type dentitions during the Albian. Except for the further specialization of the tearing- and clutching dentitions, the wide and robust cutting type dentitions present in the Albian seem to show an expansion in diet from e.g. small fish and soft-bodied animals in the Barremian to a diet of larger animals by the Albian (e.g. large fish, open marine turtles, possibly plesiosaurs). The feeding strategy used by lamniforms with this dentition to obtain their sustenance came likely both in the form of active but also through scavenging carcasses. Further investigations into lamniform disparity changes during the Early Cretaceous would require more material, most desireably from all ages of the Early Cretaceous to be analyzed in a geometric morphometric analysis. However this in

13 turn might require further field work to be carried out in order to obtain more specimens. This would be particularly advicable for pre-Albian stages (and especially the oldest stages) of the Early Cretaceous since these appear to have few lamniform specimens attributed to them in comparison to the late Early Cretaceous (in particular the Albian). Geometrical morphometric analyses should also be carried out on other organisms present during this time period to search for patterns in disparity through time, especially regarding to marine organisms. It might serve as a clue for the reason of the disparity pattern observed in the Lamniformes in this paper. When done, these might then be compared to research concerning changes in climate and environment during this time as well as provide a reason and a more complete picture of the ecomorphological changes that took place during the Early Cretaceous.

6. Acknowledgements

I would like to thank my supervisor Nicolás Campione for the guidance he has provided throughout this project. I would also like to thank my co-supervisor Benjamin Kear for making the necessary introductions. Additionally I would like to thank Mohammed Bazzi for additional guidance given concerning methodology and image design.

7. References

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7.2 Internet references

Adams, D. C., Collyer, M. L. & Sherratt, E., 2015, geomorph: Software for geometric morphometric analyses, R package version 2.1.4. http://cran.r-project.org/web/ packages/geomorph/index.html [2015-04-19] Adobe Systems Software Ireland Ltd., Adobe Photoshop CC, http://www.adobe.com/se/products/photoshop.html. [2015-04-23]. Campione, N., Quantitative Methods, http://nicolascampione.weebly.com/quantitative-methods.html [2015-04-19] Cohen, K. M., Finney, S. C., Gibbard, P. L. & Fan, J.-X., 2015, The ICS International Chronostratigraphic Chart http://www.stratigraphy.org/ICSchart/ChronostratChart2015-01.pdf. [2015-04- 21]. Inkscape, Inkscape Draw Freely https://inkscape.org/en/ [2015-04-23] Rohlf, F. J., 2015, at SUNY Stony Brook http://life.bio.sunysb.edu/morph/index.html [2015-04-22] RStudio, RStudio http://www.rstudio.com/ [2015-04-19]

8. Appendix

Figure 7. Schematic image explaining the locations of components in a as well as placement of semilandmarks in this paper.

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Table 1. Complete dataset of the 146 Early Cretaceous Lamniformes tooth specimen collected. All data was collected and entered into the dataset as found in the published material it was collected from. If a piece of information was unspecified in the litterature which it came from then information was assumed when possible, this was marked with an asterisk (*). Every row number represents one specimen each.

Row nr. Source Page Image.Label Specimen Dental.Unit 1 Cappetta 2012 232 Figure 214A UM KOB 28 lower* 2 Cappetta 2012 232 Figure 214D UM KOB 29 lower* 3 Cappetta 2012 232 Figure 214F UM KOB 30 lower* 4 Cappetta 2012 238 Figure 218A UM AUB 1 lower* 5 Cappetta 2012 239 Figure 219A UM LTX 1 lower* 6 Cappetta 2012 239 Figure 219D UM LTX 2 lower* 7 Cappetta 2012 239 Figure 219F UM ROK 1 lower 8 Cappetta 2012 240 Figure 220C UM TUI 35 lower 9 Cappetta 2012 240 Figure 220D UM TUI 37 lower 10 Cappetta 2012 240 Figure 220F UM TUI 36 lower 11 Cappetta 2012 241 Figure 221A UM KOB 14 lower* 12 Cappetta 2012 241 Figure 221D UM KOB 15 lower* 13 Cappetta 2012 241 Figure 221F UM KOB 16 lower* 14 Cappetta 2012 255 Figure 233A UM KOB 3 lower* 15 Cappetta 2012 255 Figure 233B UM KOB 4 lower* 16 Cappetta 2012 255 Figure 233E UM KOB 5 lower 17 Cappetta 2012 255 Figure 233F UM KOB 6 upper 18 Cappetta 2012 255 Figure 233H UM KOB 7 upper 19 Cappetta 2012 255 Figure 233I UM KOB 8 lower 20 Cappetta 2012 257 Figure 235A UM STO 1 lower* 21 Cappetta 2012 257 Figure 235C UM STO 2 lower* 22 Cappetta 2012 257 Figure 235F UM STO 3 lower* 23 Cappetta 2012 257 Figure 235I UM KOB 31 lower* 24 Cappetta 2012 257 Figure 235K UM KOB 32 lower* 25 Cappetta 2012 261 Figure 240A UM PER 1 lower* 26 Cappetta 2012 261 Figure 240D UM PER 2 lower* 27 Rees 2005 215 Figure 2A ZPAL P.10/8 lower* 28 Rees 2005 215 Figure 2E ZPAL P.10/10 lower* 29 Rees 2005 215 Figure 2F ZPAL P.10/9 lower* 30 Kriwet et al. 2008 282 Figure 2A MPZ 2005-4 lower* 31 Kriwet et al. 2008 282 Figure 2E MPZ 2005-5 lower* 32 Kriwet et al. 2008 282 Figure 2H MPZ 2005-6 lower* 33 Kriwet et al. 2008 282 Figure 2K MPZ 2005-7 lower* 34 Kriwet et al. 2008 282 Figure 2O MPZ 2005-8 lower* 35 Kriwet et al. 2008 282 Figure 2S MPZ 2005-9 lower* 36 Kriwet et al. 2008 282 Figure 2W MPZ 2005-10 lower* 37 Kriwet et al. 2008 283 Figure 3A MPZ 2005-11 lower* 38 Kriwet et al. 2008 283 Figure 3B MPZ 2005-12 lower* 39 Kriwet et al. 2008 283 Figure 3C MPZ 2005-13 lower* 40 Kriwet et al. 2008 283 Figure 3D MPZ 2005-14 lower*

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41 Kriwet et al. 2008 283 Figure 3F MPZ 2005-15 lower* 42 Kriwet et al. 2008 283 Figure 3G MPZ 2005-16 upper* 43 Everhart 2009 204 Figure 3D FHSM VP-17304 lower* 44 Smart 2007 377 Figure 2A NHM P 66356 lower 45 Smart 2007 377 Figure 2C NHM P 66357 lower 46 Smart 2007 377 Figure 2E NHM P 66358 lower 47 Smart 2007 378 Figure 3A NHM P 66359 upper 48 Smart 2007 378 Figure 3C NHM P 66360 upper 49 Smart 2007 378 Figure 3E NHM P 66361 upper 50 Siverson 2007 et al. 943 Text-fig. 3A SMU 76283 lower* 51 Siverson 2007 et al. 943 Text-fig. 3C SMU 76282 lower* 52 Siverson 2007 et al. 943 Text-fig. 3F SMU 76284 lower* 53 Siverson 2007 et al. 943 Text-fig. 3G SMU 76285 lower* 54 Siverson 2007 et al. 943 Text-fig. 3J SMU 76286 lower* 55 Siverson 2007 et al. 943 Text-fig. 3L SMU 76287 lower* 56 Siverson 2007 et al. 943 Text-fig. 3N SMU 76288 lower* 57 Siverson 2007 et al. 943 Text-fig. 3P SMU 76289 lower* 58 Siverson 2007 et al. 944 Text-fig. 4A SMU 76311 lower* 59 Siverson 2007 et al. 947 Plate 1, figure 1 SMU 76313 lower* 60 Siverson 2007 et al. 947 Plate 1, figure 4 SMU 76314 lower* 61 Siverson 2007 et al. 947 Plate 1, figure 5 SMU 76315 lower* 62 Siverson 2007 et al. 947 Plate 1, figure 8 SMU 76312 lower* 63 Siverson 2007 et al. 947 Plate 1, figure 10 SMU 76316 lower* 64 Siverson 2007 et al. 947 Plate 1, figure 12 SMU 76317 lower* 65 Siverson 2007 et al. 947 Plate 1, figure 13 SMU 76318 lower* 66 Siverson 2007 et al. 947 Plate 1, figure 16 SMU 76319 lower* 67 Siverson 2007 et al. 947 Plate 1, figure 17 SMU 76320 lower* 68 Siverson 2007 et al. 947 Plate 1, figure 20 SMU 76321 lower* 69 Siverson 2007 et al. 947 Plate 1, figure 21 SMU 76322 lower* 70 Siverson 2007 et al. 947 Plate 1, figure 23 SMU 76323 lower* 71 Siverson 2007 et al. 947 Plate 1, figure 26 SMU 76324 lower* 72 Kriwet 2006 540 Figure 2A BMNH P.36287 lower 73 Kriwet 2006 540 Figure 2B BMNH P.36288 lower* 74 Kriwet 2006 540 Figure 3A BMNH P.36289 lower* 75 Kriwet 2006 540 Figure 3D BMNH P.36290 lower* 76 Kriwet 2006 540 Figure 3H BMNH P.36291 lower* 77 Kriwet 2006 540 Figure 3G lower* 78 Kriwet 2006 540 Figure 3J lower* 79 Shimada 2007 513 Figure 2J lower* 80 Siverson et al. 2013 92 Fig. 4D NHMUK PV P73050 upper 81 Siverson et al. 2013 92 Fig. 4H NHMUK PV P73051 upper 82 Siverson et al. 2013 92 Fig. 4L NHMUK PV P73052 upper 83 Siverson et al. 2013 92 Fig. 4M NHMUK PV P73053 upper 84 Siverson et al. 2013 92 Fig. 4S NHMUK PV P73054 upper 85 Siverson et al. 2013 92 Fig. 4U NHMUK PV P73055 upper 86 Siverson et al. 2013 92 Fig. 4B' NHMUK PV P73056 upper

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87 Siverson et al. 2013 92 Fig. 4C' NHMUK PV P73057 upper 88 Siverson et al. 2013 92 Fig. 4F' NHMUK PV P73058 upper 89 Siverson et al. 2013 94 Fig. 5A NHMUK PV P73059 upper 90 Siverson et al. 2013 94 Fig. 5E NHMUK PV P73060 upper 91 Siverson et al. 2013 94 Fig. 5G NHMUK PV P73061 upper 92 Siverson et al. 2013 94 Fig. 5J NHMUK PV P73062 upper 93 Siverson et al. 2013 94 Fig. 5O NHMUK PV P73063 upper 94 Siverson et al. 2013 94 Fig. 5Q NHMUK PV P73064 upper 95 Siverson et al. 2013 94 Fig. 5U NHMUK PV P73065 upper 96 Siverson et al. 2013 94 Fig. 5Y NHMUK PV P73066 upper 97 Siverson et al. 2013 94 Fig. 5C' NHMUK PV P73067 upper 98 Siverson et al. 2013 94 Fig. 5F' NHMUK PV P73068 upper 99 Siverson et al. 2013 95 Fig. 6A NHMUK PV P73069 lower 100 Siverson et al. 2013 95 Fig. 6E NHMUK PV P73070 lower 101 Siverson et al. 2013 95 Fig. 6L NHMUK PV P73071 lower 102 Siverson et al. 2013 95 Fig. 6O NHMUK PV P73072 lower 103 Siverson et al. 2013 96 Fig. 7C NHMUK PV P73073 lower 104 Siverson et al. 2013 96 Fig. 7H NHMUK PV P73074 lower 105 Siverson et al. 2013 96 Fig. 7I NHMUK PV P73075 lower 106 Siverson et al. 2013 96 Fig. 7M NHMUK PV P73076 lower 107 Siverson et al. 2013 96 Fig. 7T NHMUK PV P73077 lower 108 Siverson et al. 2013 96 Fig. 7W NHMUK PV P73078 lower 109 Siverson et al. 2013 96 Fig. 7Y NHMUK PV P73079 lower 110 Siverson et al. 2013 96 Fig. 7C' NHMUK PV P73080 lower 111 Siverson et al. 2013 96 Fig. 7G' NHMUK PV P73081 lower 112 Siverson et al. 2013 98 Fig. 9 (tooth LP3) NHMUK PV P73082 upper* 113 Siverson et al. 2013 98 Fig. 9 (tooth LP9) NHMUK PV P73083 upper* 114 Siverson et al. 2013 97 Fig. 8D SMU 76857 upper* 115 Siverson et al. 2013 97 Fig. 8F SMU 76858 upper* 116 Siverson et al. 2013 97 Fig. 8H SMU 76859 upper* 117 Siverson et al. 2013 97 Fig. 8L SMU 76860 lower* 118 Siverson et al. 2013 97 Fig. 8R SMU 76861 lower* 119 Kriwet et al. 2009 321 Figure 4J MPZ 2005-4 lower* 120 Kriwet et al. 2009 321 Figure 4N MPZ 2005-4 lower* 121 Kriwet et al. 2009 321 Figure 4R MPZ 2005-4 lower* 122 Schmitz et al. 2010 286 Fig. 3.1 NLH 102.973 lower* 123 Schmitz et al. 2010 286 Fig. 3.3 NLH 102.974 lower* 124 Schmitz et al. 2010 286 Fig. 3.6 NLH 102.976 lower* 125 Schmitz et al. 2010 286 Fig. 3.9 NLH 102.977 lower* 126 Schmitz et al. 2010 286 Fig. 3.11 NLH 102.978 lower* 127 Schmitz et al. 2010 286 Fig. 3.13 NLH 102.979 upper (?) 128 Cuny et al. 2010 618 Fig. 2.5 OED5 lower* 129 Schlüter and Schwarzhans 1978 72 Tafel 2, Fig. 1 upper* 130 Benton et al. 2000 242 Fig. 13B lower* 131 Benton et al. 2000 242 Fig. 13D lower* 132 Siverson 1997 462 Figure 4A WAM 96.2.42 upper

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133 Siverson 1997 462 Figure 4F WAM 96.2.43 upper 134 Siverson 1997 462 Figure 4H WAM 96.2.44 lower? 135 Siverson 1997 462 Figure 4J WAM 96.2.66 lower* 136 Siverson 1997 462 Figure 4L WAM 96.2.74 upper? 137 Siverson 1997 462 Figure 4O WAM 96.2.75 lower 138 Siverson 1997 462 Figure 4R WAM 96.2.80 lower* 139 Siverson 1997 462 Figure 4T WAM 96.2.81 lower 140 Siverson 1997 462 Figure 4W WAM 96.2.82 lower 141 Siverson 1997 462 Figure 4X WAM 96.2.85 lower* 142 Antunes and Cappetta 2002 159 Planche 7, Fig. 1a ANG 51 lower 143 Antunes and Cappetta 2002 165 Planche 10, Fig. 3a ANG 93 lower* 144 Antunes and Cappetta 2002 167 Planche 11, Fig. 10 ANG 119 lower 145 Bouaziz et al. 1988 337 Fig. 2K2 lower* 146 Bouaziz et al. 1988 337 Fig. 2L1 lower*

(continued) Row nr. Side (L=Left, R=Right side Relative.Position View Order of jaw) 1 anterior labial Lamniformes 2 lateral labial Lamniformes 3 anterolateral labial Lamniformes 4 anterolateral labial Lamniformes 5 anterior labial Lamniformes 6 anterior labial Lamniformes 7 lateral labial Lamniformes 8 anterior labial Lamniformes 9 anterolateral lingual Lamniformes 10 lateral labial Lamniformes 11 anterior labial Lamniformes 12 anterolateral labial Lamniformes 13 anterior labial Lamniformes 14 parasymphyseal labial Lamniformes 15 anterior labial Lamniformes 16 anterolateral labial Lamniformes 17 labial Lamniformes 18 very lateral lingual Lamniformes 19 very lateral lingual Lamniformes 20 anterior labial Lamniformes 21 anterolateral labial Lamniformes 22 anterolateral labial Lamniformes 23 anterior labial Lamniformes 24 anterolateral labial Lamniformes 25 anterior labial Lamniformes 26 anterolateral labial Lamniformes 27 anterior labial Lamniformes 28 anterior labial Lamniformes 29 labial Lamniformes

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30 anterolateral labial Lamniformes 31 symphyseal? labial Lamniformes 32 anterior labial Lamniformes 33 lateral labial Lamniformes 34 lateral labial Lamniformes 35 lateral labial Lamniformes 36 posterior labial Lamniformes 37 anterolateral labial Lamniformes 38 lateral labial Lamniformes 39 lateral lingual Lamniformes 40 lateral lingual Lamniformes 41 anterolateral labial Lamniformes 42 intermediate? labial Lamniformes 43 Lamniformes 44 L lateral labial Lamniformes 45 R lateral labial Lamniformes 46 L lateral labial Lamniformes 47 L anterior labial Lamniformes 48 L lateral labial Lamniformes 49 L lateral labial Lamniformes 50 anterolateral labial Lamniformes 51 lateral labial Lamniformes 52 lateral labial Lamniformes 53 lateral labial Lamniformes 54 lateral labial Lamniformes 55 lateroposterior labial Lamniformes 56 lateral labial Lamniformes 57 posterior labial Lamniformes 58 lateral labial Lamniformes 59 anterior labial Lamniformes 60 anterolateral labial Lamniformes 61 lateral labial Lamniformes 62 lateral labial Lamniformes 63 lateroposterior labial Lamniformes 64 lateral labial Lamniformes 65 lateral labial Lamniformes 66 lateroposterior labial Lamniformes 67 lateral labial Lamniformes 68 lateroposterior labial Lamniformes 69 lateroposterior labial Lamniformes 70 posterior labial Lamniformes 71 posterior labial Lamniformes 72 lateral lingual Lamniformes 73 anterior labial Lamniformes 74 anterior? labial Lamniformes 75 lateral labial Lamniformes

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76 lateroposterior labial Lamniformes 77 lateral lingual Lamniformes 78 lateroposterior lingual Lamniformes 79 labial Lamniformes 80 R anterior labial Lamniformes 81 L parasymphyseal labial Lamniformes 82 R? anterior labial Lamniformes 83 parasymphyseal labial Lamniformes 84 R anterior labial Lamniformes 85 R anterior labial Lamniformes 86 R anterior labial Lamniformes 87 R anterior labial Lamniformes 88 L anterior labial Lamniformes 89 R lateroposterior labial Lamniformes 90 lateroposterior labial Lamniformes 91 L lateroposterior labial Lamniformes 92 L lateroposterior labial Lamniformes 93 L lateroposterior labial Lamniformes 94 L lateroposterior labial Lamniformes 95 L lateroposterior labial Lamniformes 96 R lateroposterior labial Lamniformes 97 R lateroposterior labial Lamniformes 98 R lateroposterior labial Lamniformes 99 L anterior labial Lamniformes 100 L anterior labial Lamniformes 101 R anterior labial Lamniformes 102 R anterior labial Lamniformes 103 R lateroposterior labial Lamniformes 104 R lateroposterior labial Lamniformes 105 L lateroposterior labial Lamniformes 106 L lateroposterior labial Lamniformes 107 L lateroposterior labial Lamniformes 108 L lateroposterior labial Lamniformes 109 L lateroposterior labial Lamniformes 110 L lateroposterior labial Lamniformes 111 L lateroposterior labial Lamniformes 112 lateroposterior labial Lamniformes 113 lateroposterior labial Lamniformes 114 L lateroposterior labial Lamniformes 115 L lateroposterior labial Lamniformes 116 L lateroposterior labial Lamniformes 117 L lateroposterior labial Lamniformes 118 R lateroposterior labial Lamniformes 119 anterolateral labial Lamniformes 120 lateral labial Lamniformes 121 lateral labial Lamniformes

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122 anterior labial Lamniformes 123 anterior labial Lamniformes 124 anterior labial Lamniformes 125 lateral labial Lamniformes 126 posterolateral labial Lamniformes 127 parasymphyseal labial Lamniformes 128 labial Lamniformes 129 labial Lamniformes 130 labial Lamniformes* 131 labial Lamniformes* 132 R anterior labial Lamniformes 133 L lateral labial Lamniformes 134 R? anterior labial Lamniformes 135 labial Lamniformes 136 R? lateral labial Lamniformes 137 L lateral labial Lamniformes 138 anterior labial Lamniformes 139 L anterior labial Lamniformes 140 R anterior labial Lamniformes 141 lateral labial Lamniformes 142 anterior labial Lamniformes 143 anterior labial Lamniformes 144 anteriolateral labial Lamniformes 145 labial Lamniformes 146 labial Lamniformes

(continued) Row nr. Family Genus Species Continent Country 1 Cardabiodontidae Pseudoisurus aff. tomosus Asia (central) Kazakhstan 2 Cardabiodontidae Pseudoisurus aff. tomosus Asia (central) Kazakhstan 3 Cardabiodontidae Pseudoisurus aff. tomosus Asia (central) Kazakhstan 4 Paraisuridae Paraisurus macrorhiza Europe France 5 Pseudoscapanorhynchidae Leptostyrax bicuspidatus North America USA 6 Pseudoscapanorhynchidae Leptostyrax bicuspidatus North America USA 7 Pseudoscapanorhynchidae Leptostyrax bicuspidatus North America USA 8 Pseudoscapanorhynchidae Protolamna sokolovi Europe France 9 Pseudoscapanorhynchidae Protolamna sokolovi Europe France 10 Pseudoscapanorhynchidae Protolamna sokolovi Europe France 11 Pseudoscapanorhynchidae Pseudoscarpanorhynchus aff. compressidens Asia (central) Kazakhstan 12 Pseudoscapanorhynchidae Pseudoscarpanorhynchus aff. compressidens Asia (central) Kazakhstan 13 Pseudoscapanorhynchidae Pseudoscarpanorhynchus aff. compressidens Asia (central) Kazakhstan 14 Cretodus semiplicatus Asia (central) Kazakhstan 15 Cretodus semiplicatus Asia (central) Kazakhstan 16 Cretodus semiplicatus Asia (central) Kazakhstan 17 Cretodus semiplicatus Asia (central) Kazakhstan 18 Cretodus semiplicatus Asia (central) Kazakhstan 19 Cretodus semiplicatus Asia (central) Kazakhstan

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20 Dwardius siversoni Europe Russia 21 Dwardius siversoni Europe Russia 22 Dwardius siversoni Europe Russia 23 Dwardius sp. Asia (central) Kazakhstan 24 Dwardius sp. Asia (central) Kazakhstan 25 Priscusurus adruptodontus South America Peru 26 Priscusurus adruptodontus South America Peru 27 Cretoxyrhinidae Protolamna sp. Europe Poland 28 Cretoxyrhinidae Protolamna sp. Europe Poland 29 Cretoxyrhinidae Protolamna sp. Europe Poland 30 Eoptolamnidae Eoptolamna eccentrolopha Europe Spain 31 Eoptolamnidae Eoptolamna eccentrolopha Europe Spain 32 Eoptolamnidae Eoptolamna eccentrolopha Europe Spain 33 Eoptolamnidae Eoptolamna eccentrolopha Europe Spain 34 Eoptolamnidae Eoptolamna eccentrolopha Europe Spain 35 Eoptolamnidae Eoptolamna eccentrolopha Europe Spain 36 Eoptolamnidae Eoptolamna eccentrolopha Europe Spain 37 Eoptolamnidae Eoptolamna eccentrolopha Europe Spain 38 Eoptolamnidae Eoptolamna eccentrolopha Europe Spain 39 Eoptolamnidae Eoptolamna eccentrolopha Europe Spain 40 Eoptolamnidae Eoptolamna eccentrolopha Europe Spain 41 Eoptolamnidae Eoptolamna eccentrolopha Europe Spain 42 Eoptolamnidae Eoptolamna eccentrolopha Europe Spain 43 Odontaspididae Carcharias amonensis North America USA 44 Squalicorax primaevus Europe England 45 Anacoracidae Squalicorax primaevus Europe England 46 Anacoracidae Squalicorax primaevus Europe England 47 Anacoracidae Squalicorax primaevus Europe England 48 Anacoracidae Squalicorax primaevus Europe England 49 Anacoracidae Squalicorax primaevus Europe England 50 Anacoracidae Squalicorax priscoserratus sp. nov. North America USA 51 Anacoracidae Squalicorax priscoserratus sp. nov. North America USA 52 Anacoracidae Squalicorax priscoserratus sp. nov. North America USA 53 Anacoracidae Squalicorax priscoserratus sp. nov. North America USA 54 Anacoracidae Squalicorax priscoserratus sp. nov. North America USA 55 Anacoracidae Squalicorax priscoserratus sp. nov. North America USA 56 Anacoracidae Squalicorax priscoserratus sp. nov. North America USA 57 Anacoracidae Squalicorax priscoserratus sp. nov. North America USA 58 Anacoracidae Squalicorax aff. S. baharijensis North America USA 59 Anacoracidae Squalicorax pawpawensis sp. nov. North America USA 60 Anacoracidae Squalicorax pawpawensis sp. nov. North America USA 61 Anacoracidae Squalicorax pawpawensis sp. nov. North America USA 62 Anacoracidae Squalicorax pawpawensis sp. nov. North America USA 63 Anacoracidae Squalicorax pawpawensis sp. nov. North America USA 64 Anacoracidae Squalicorax pawpawensis sp. nov. North America USA 65 Anacoracidae Squalicorax pawpawensis sp. nov. North America USA

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66 Anacoracidae Squalicorax pawpawensis sp. nov. North America USA 67 Anacoracidae Squalicorax pawpawensis sp. nov. North America USA 68 Anacoracidae Squalicorax pawpawensis sp. nov. North America USA 69 Anacoracidae Squalicorax pawpawensis sp. nov. North America USA 70 Anacoracidae Squalicorax pawpawensis sp. nov. North America USA 71 Anacoracidae Squalicorax pawpawensis sp. nov. North America USA 72 incertae sedis Priscusurus adruptodontus sp. nov. South America Peru 73 incertae sedis Priscusurus adruptodontus sp. nov. South America Peru 74 incertae sedis Priscusurus adruptodontus sp. nov. South America Peru 75 incertae sedis Priscusurus adruptodontus sp. nov. South America Peru 76 incertae sedis Priscusurus adruptodontus sp. nov. South America Peru 77 incertae sedis Priscusurus adruptodontus sp. nov. South America Peru 78 incertae sedis Priscusurus adruptodontus sp. nov. South America Peru 79 Odontaspididae cf. Johnlongia sp. Europe England 80 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 81 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 82 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 83 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 84 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 85 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 86 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 87 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 88 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 89 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 90 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 91 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 92 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 93 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 94 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 95 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 96 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 97 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 98 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 99 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 100 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 101 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 102 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 103 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 104 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 105 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 106 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 107 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 108 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 109 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 110 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 111 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan

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112 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 113 Cretoxyrhinidae Cretoxyrhina vraconensis Asia (central) Kazakhstan 114 Cretoxyrhinidae Cretoxyrhina vraconensis North America USA 115 Cretoxyrhinidae Cretoxyrhina vraconensis North America USA 116 Cretoxyrhinidae Cretoxyrhina vraconensis North America USA 117 Cretoxyrhinidae Cretoxyrhina vraconensis North America USA 118 Cretoxyrhinidae Cretoxyrhina vraconensis North America USA 119 incertae sedis Europe Spain 120 incertae sedis Europe Spain 121 incertae sedis Europe Spain 122 Eoptolamnidae Leptostyrax stychi sp. nov. Europe Germany 123 Eoptolamnidae Leptostyrax stychi sp. nov. Europe Germany 124 Eoptolamnidae Protolamna sarstedtensis sp. nov. Europe Germany 125 Eoptolamnidae Protolamna sarstedtensis sp. nov. Europe Germany 126 Eoptolamnidae Protolamna sarstedtensis sp. nov. Europe Germany 127 Eoptolamnidae Protolamna sarstedtensis sp. nov. Europe Germany 128 Cretoxyrhinidae ?Cretodus Africa Tunisia 129 sp. Africa Tunisia 130 Cretodus Africa Tunisia 131 Protolamna sp. Africa Tunisia 132 Cretoxyrhinidae Archaeolamna sp. Australia Australia 133 Cretoxyrhinidae Archaeolamna sp. Australia Australia 134 Cretoxyrhinidae Archaeolamna sp. Australia Australia 135 Cretoxyrhinidae Leptostyrax sp. Austalia Australia 136 Cretoxyrhinidae incertae sedis Australia Australia 137 Cretoxyrhinidae Cretolamna sp. Australia Australia 138 Cretoxyrhinidae Paraisurus aff. compressus Australia Australia 139 Odontaspididae Carcharias striatula Australia Australia 140 Odontaspididae Carcharias striatula Australia Australia 141 Anacoracidae Squalicorax primaevus Australia Australia 142 Anacoracidae Squalicorax sp. Africa Angola 143 Cretoxyrhinidae Leptostyrax macrorhiza Africa Angola 144 Cretoxyrhinidae Protolamna sp. Africa Angola 145 Cretoxyrhinidae Cretodus ? Africa Tunisia 146 Cretoxyrhinidae Protolamna sp. Africa Tunisia

(continued) Row nr. Locale 1 Kolbay 2 Kolbay 3 Kolbay 4 Aube, Paris basin 5 Lake Texoma, Texas 6 Lake Texoma, Texas 7 Roanoke, Texas 8 La Tuilière, Apt region, Vaucluse 9 La Tuilière, Apt region, Vaucluse

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10 La Tuilière, Apt region, Vaucluse 11 Kolbay 12 Kolbay 13 Kolbay 14 Kolbay 15 Kolbay 16 Kolbay 17 Kolbay 18 Kolbay 19 Kolbay 20 Stojlenski quarry, Stary Oskol, near Belgorod. 21 Stojlenski quarry, Stary Oskol, near Belgorod. 22 Stojlenski quarry, Stary Oskol, near Belgorod. 23 Kolbay 24 Kolbay 25 26 27 Clay pit in Wawał 28 Clay pit in Wawał 29 Clay pit in Wawał 30 Vallipón in NW part of Maestrat sub-basin near the city of Castellote (approximately 150 km SE of Zaragoza) 31 Vallipón in NW part of Maestrat sub-basin near the city of Castellote (approximately 150 km SE of Zaragoza) 32 Vallipón in NW part of Maestrat sub-basin near the city of Castellote (approximately 150 km SE of Zaragoza) 33 Vallipón in NW part of Maestrat sub-basin near the city of Castellote (approximately 150 km SE of Zaragoza) 34 Vallipón in NW part of Maestrat sub-basin near the city of Castellote (approximately 150 km SE of Zaragoza) 35 Vallipón in NW part of Maestrat sub-basin near the city of Castellote (approximately 150 km SE of Zaragoza) 36 Vallipón in NW part of Maestrat sub-basin near the city of Castellote (approximately 150 km SE of Zaragoza) 37 Vallipón in NW part of Maestrat sub-basin near the city of Castellote (approximately 150 km SE of Zaragoza) 38 Vallipón in NW part of Maestrat sub-basin near the city of Castellote (approximately 150 km SE of Zaragoza) 39 Vallipón in NW part of Maestrat sub-basin near the city of Castellote (approximately 150 km SE of Zaragoza) 40 Vallipón in NW part of Maestrat sub-basin near the city of Castellote (approximately 150 km SE of Zaragoza) 41 Vallipón in NW part of Maestrat sub-basin near the city of Castellote (approximately 150 km SE of Zaragoza) 42 Vallipón in NW part of Maestrat sub-basin near the city of Castellote (approximately 150 km SE of Zaragoza) 43 Champion shell bed, Kansas 44 Gault clays of Folkestone, NE of Leighton Buzzard 45 Gault clays of Folkestone, NE of Leighton Buzzard 46 Gault clays of Folkestone, NE of Leighton Buzzard 47 Gault clays of Folkestone, NE of Leighton Buzzard 48 Gault clays of Folkestone, NE of Leighton Buzzard 49 Gault clays of Folkestone, NE of Leighton Buzzard 50 Motorola locality, Pawpaw shale, Texas 51 Motorola locality, Pawpaw shale, Texas 52 Motorola locality, Pawpaw shale, Texas 53 Motorola locality, Pawpaw shale, Texas 54 Motorola locality, Pawpaw shale, Texas 55 Motorola locality, Pawpaw shale, Texas

28

56 Motorola locality, Pawpaw shale, Texas 57 Motorola locality, Pawpaw shale, Texas 58 Motorola locality, Pawpaw shale, Texas 59 Motorola locality, Pawpaw shale, Texas 60 Motorola locality, Pawpaw shale, Texas 61 Motorola locality, Pawpaw shale, Texas 62 Motorola locality, Pawpaw shale, Texas 63 Motorola locality, Pawpaw shale, Texas 64 Motorola locality, Pawpaw shale, Texas 65 Motorola locality, Pawpaw shale, Texas 66 Motorola locality, Pawpaw shale, Texas 67 Motorola locality, Pawpaw shale, Texas 68 Motorola locality, Pawpaw shale, Texas 69 Motorola locality, Pawpaw shale, Texas 70 Motorola locality, Pawpaw shale, Texas 71 Motorola locality, Pawpaw shale, Texas 72 Locality 3, Ammonite Valley, Amotape-Tahuin Massif, 27 miles east of Lobitos 73 Locality 3, Ammonite Valley, Amotape-Tahuin Massif, 27 miles east of Lobitos 74 Locality 3, Ammonite Valley, Amotape-Tahuin Massif, 27 miles east of Lobitos 75 Locality 3, Ammonite Valley, Amotape-Tahuin Massif, 27 miles east of Lobitos 76 Locality 3, Ammonite Valley, Amotape-Tahuin Massif, 27 miles east of Lobitos 77 Locality 3, Ammonite Valley, Amotape-Tahuin Massif, 27 miles east of Lobitos 78 Locality 3, Ammonite Valley, Amotape-Tahuin Massif, 27 miles east of Lobitos 79 80 Kolbay, Mangysklak 81 Kolbay, Mangysklak 82 Kolbay, Mangysklak 83 Kolbay, Mangysklak 84 Kolbay, Mangysklak 85 Kolbay, Mangysklak 86 Kolbay, Mangysklak 87 Kolbay, Mangysklak 88 Kolbay, Mangysklak 89 Kolbay, Mangysklak 90 Kolbay, Mangysklak 91 Kolbay, Mangysklak 92 Kolbay, Mangysklak 93 Kolbay, Mangysklak 94 Kolbay, Mangysklak 95 Kolbay, Mangysklak 96 Kolbay, Mangysklak 97 Kolbay, Mangysklak 98 Kolbay, Mangysklak 99 Kolbay, Mangysklak 100 Kolbay, Mangysklak 101 Kolbay, Mangysklak

29

102 Kolbay, Mangysklak 103 Kolbay, Mangysklak 104 Kolbay, Mangysklak 105 Kolbay, Mangysklak 106 Kolbay, Mangysklak 107 Kolbay, Mangysklak 108 Kolbay, Mangysklak 109 Kolbay, Mangysklak 110 Kolbay, Mangysklak 111 Kolbay, Mangysklak 112 Kolbay, Mangysklak 113 Kolbay, Mangysklak 114 Motorola site, Mortoniceras rostratum Zone, Pawpaw shale, Texas 115 Motorola site, Mortoniceras rostratum Zone, Pawpaw shale, Texas 116 Motorola site, Mortoniceras rostratum Zone, Pawpaw shale, Texas 117 Motorola site, Mortoniceras rostratum Zone, Pawpaw shale, Texas 118 Motorola site, Mortoniceras rostratum Zone, Pawpaw shale, Texas 119 Oliete subbasin, Iberian basin, NE-Spain 120 Oliete subbasin, Iberian basin, NE-Spain 121 Oliete subbasin, Iberian basin, NE-Spain 122 'Gott'' Clay pit approx. 20 km SE of Hannover, near the city of Sarstedt, northern Germany 123 'Gott'' Clay pit approx. 20 km SE of Hannover, near the city of Sarstedt, northern Germany 124 'Gott'' Clay pit approx. 20 km SE of Hannover, near the city of Sarstedt, northern Germany 125 'Gott'' Clay pit approx. 20 km SE of Hannover, near the city of Sarstedt, northern Germany 126 'Gott'' Clay pit approx. 20 km SE of Hannover, near the city of Sarstedt, northern Germany 127 'Gott'' Clay pit approx. 20 km SE of Hannover, near the city of Sarstedt, northern Germany 128 Oum ed Diab 129 Near Ksar Krerachfa, south of the city Medenine, south Tunisia 130 Tataouine region, southern Tunisia 131 Tataouine region, southern Tunisia 132 Black Dam, lower part of Gearle Siltstone, Giralia Anticline, Southern Carnarvon basin, 133 Black Dam, lower part of Gearle Siltstone, Giralia Anticline, Southern Carnarvon basin, Western Australia 134 Black Dam, lower part of Gearle Siltstone, Giralia Anticline, Southern Carnarvon basin, Western Australia 135 Black Dam, lower part of Gearle Siltstone, Giralia Anticline, Southern Carnarvon basin, Western Australia 136 Black Dam, lower part of Gearle Siltstone, Giralia Anticline, Southern Carnarvon basin, Western Australia 137 Black Dam, lower part of Gearle Siltstone, Giralia Anticline, Southern Carnarvon basin, Western Australia 138 Black Dam, lower part of Gearle Siltstone, Giralia Anticline, Southern Carnarvon basin, Western Australia 139 Black Dam, lower part of Gearle Siltstone, Giralia Anticline, Southern Carnarvon basin, Western Australia 140 Black Dam, lower part of Gearle Siltstone, Giralia Anticline, Southern Carnarvon basin, Western Australia 141 Black Dam, lower part of Gearle Siltstone, Giralia Anticline, Southern Carnarvon basin, Western Australia 142 Locality 19, Cubal da Hanha, 19 km NE of Lobito, Benguela basin 143 Locality 19, Cubal da Hanha, 19 km NE of Lobito, Benguela basin 144 Locality 19, Cubal da Hanha, 19 km NE of Lobito, Benguela basin 145 Locality RH 45, Foum Tatahouine region, southern Tunisia 146 Locality RH 45, Foum Tatahouine region, southern Tunisia

(continued) 30

Row nr. Formation Era Period Epoch Sub Age 1 Mesozoic Cretaceous Lower late Albian 2 Mesozoic Cretaceous Lower late Albian 3 Mesozoic Cretaceous Lower late Albian 4 Mesozoic Cretaceous Lower Albian 5 Mesozoic Cretaceous Lower Albian 6 Mesozoic Cretaceous Lower Albian 7 Mesozoic Cretaceous Lower Albian 8 Mesozoic Cretaceous Lower middle to late Albian 9 Mesozoic Cretaceous Lower middle to late Albian 10 Mesozoic Cretaceous Lower middle to late Albian 11 Mesozoic Cretaceous Lower late Albian 12 Mesozoic Cretaceous Lower late Albian 13 Mesozoic Cretaceous Lower late Albian 14 Mesozoic Cretaceous Lower late Albian 15 Mesozoic Cretaceous Lower late Albian 16 Mesozoic Cretaceous Lower late Albian 17 Mesozoic Cretaceous Lower late Albian 18 Mesozoic Cretaceous Lower late Albian 19 Mesozoic Cretaceous Lower late Albian 20 Mesozoic Cretaceous Lower Albian 21 Mesozoic Cretaceous Lower Albian 22 Mesozoic Cretaceous Lower Albian 23 Mesozoic Cretaceous Lower late Albian 24 Mesozoic Cretaceous Lower late Albian 25 Muerto Mesozoic Cretaceous Lower middle Albian formation 26 Muerto Limestone Mesozoic Cretaceous Lower middle Albian formation 27 Mesozoic Cretaceous Lower late Valanginian 28 Mesozoic Cretaceous Lower late Valanginian 29 Mesozoic Cretaceous Lower late Valanginian 30 Artoles formation Mesozoic Cretaceous Lower late Barremian 31 Artoles formation Mesozoic Cretaceous Lower late Barremian 32 Artoles formation Mesozoic Cretaceous Lower late Barremian 33 Artoles formation Mesozoic Cretaceous Lower late Barremian 34 Artoles formation Mesozoic Cretaceous Lower late Barremian 35 Artoles formation Mesozoic Cretaceous Lower late Barremian 36 Artoles formation Mesozoic Cretaceous Lower late Barremian 37 Artoles formation Mesozoic Cretaceous Lower late Barremian 38 Artoles formation Mesozoic Cretaceous Lower late Barremian 39 Artoles formation Mesozoic Cretaceous Lower late Barremian 40 Artoles formation Mesozoic Cretaceous Lower late Barremian 41 Artoles formation Mesozoic Cretaceous Lower late Barremian 42 Artoles formation Mesozoic Cretaceous Lower late Barremian 43 Kiowa formation Mesozoic Cretaceous Lower late Albian 44 Mesozoic Cretaceous Lower middle Albian 45 Mesozoic Cretaceous Lower middle Albian

31

46 Mesozoic Cretaceous Lower upper Albian 47 Mesozoic Cretaceous Lower middle Albian 48 Mesozoic Cretaceous Lower upper Albian 49 Mesozoic Cretaceous Lower middle Albian 50 Pawpaw formation Mesozoic Cretaceous Lower late Albian 51 Pawpaw formation Mesozoic Cretaceous Lower late Albian 52 Pawpaw formation Mesozoic Cretaceous Lower late Albian 53 Pawpaw formation Mesozoic Cretaceous Lower late Albian 54 Pawpaw formation Mesozoic Cretaceous Lower late Albian 55 Pawpaw formation Mesozoic Cretaceous Lower late Albian 56 Pawpaw formation Mesozoic Cretaceous Lower late Albian 57 Pawpaw formation Mesozoic Cretaceous Lower late Albian 58 Pawpaw formation Mesozoic Cretaceous Lower late Albian 59 Pawpaw formation Mesozoic Cretaceous Lower late Albian 60 Pawpaw formation Mesozoic Cretaceous Lower late Albian 61 Pawpaw formation Mesozoic Cretaceous Lower late Albian 62 Pawpaw formation Mesozoic Cretaceous Lower late Albian 63 Pawpaw formation Mesozoic Cretaceous Lower late Albian 64 Pawpaw formation Mesozoic Cretaceous Lower late Albian 65 Pawpaw formation Mesozoic Cretaceous Lower late Albian 66 Pawpaw formation Mesozoic Cretaceous Lower late Albian 67 Pawpaw formation Mesozoic Cretaceous Lower late Albian 68 Pawpaw formation Mesozoic Cretaceous Lower late Albian 69 Pawpaw formation Mesozoic Cretaceous Lower late Albian 70 Pawpaw formation Mesozoic Cretaceous Lower late Albian 71 Pawpaw formation Mesozoic Cretaceous Lower late Albian 72 Muerto Limestone Mesozoic Cretaceous Lower middle? Albian formation 73 Muerto Limestone Mesozoic Cretaceous Lower middle? Albian formation 74 Muerto Limestone Mesozoic Cretaceous Lower middle? Albian formation 75 Muerto Limestone Mesozoic Cretaceous Lower middle? Albian formation 76 Muerto Limestone Mesozoic Cretaceous Lower middle? Albian formation 77 Muerto Limestone Mesozoic Cretaceous Lower middle? Albian formation 78 Muerto Limestone Mesozoic Cretaceous Lower middle? Albian formation 79 Mesozoic Cretaceous Lower early Aptian 80 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 81 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 82 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 83 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 84 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 85 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 86 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 87 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 88 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 89 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 90 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian

32

91 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 92 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 93 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 94 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 95 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 96 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 97 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 98 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 99 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 100 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 101 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 102 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 103 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 104 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 105 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 106 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 107 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 108 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 109 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 110 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 111 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 112 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 113 Mesozoic Cretaceous Lower latest late/earliest early Albian/Cenomanian 114 Pawpaw formation Mesozoic Cretaceous Lower late Albian 115 Pawpaw formation Mesozoic Cretaceous Lower late Albian 116 Pawpaw formation Mesozoic Cretaceous Lower late Albian 117 Pawpaw formation Mesozoic Cretaceous Lower late Albian 118 Pawpaw formation Mesozoic Cretaceous Lower late Albian 119 upper Blesa formation Mesozoic Cretaceous Lower late Barremian 120 upper Blesa formation Mesozoic Cretaceous Lower late Barremian 121 upper Blesa formation Mesozoic Cretaceous Lower late Barremian 122 Mesozoic Cretaceous Lower early Barremian 123 Mesozoic Cretaceous Lower early Barremian 124 Mesozoic Cretaceous Lower early Barremian 125 Mesozoic Cretaceous Lower early Barremian 126 Mesozoic Cretaceous Lower early Barremian 127 Mesozoic Cretaceous Lower early Barremian 128 Aïn el Guettar Mesozoic Cretaceous Lower early Albian formation, Oum ed Diab group 129 Mesozoic Cretaceous Lower 130 Chenini formation Mesozoic Cretaceous Lower early Albian 131 Chenini formation Mesozoic Cretaceous Lower early Albian 132 Mesozoic Cretaceous Lower middle to late Albian 133 Mesozoic Cretaceous Lower middle to late Albian 134 Mesozoic Cretaceous Lower middle to late Albian 135 Mesozoic Cretaceous Lower middle to late Albian

33

136 Mesozoic Cretaceous Lower middle to late Albian 137 Mesozoic Cretaceous Lower middle to late Albian 138 Mesozoic Cretaceous Lower middle to late Albian 139 Mesozoic Cretaceous Lower middle to late Albian 140 Mesozoic Cretaceous Lower middle to late Albian 141 Mesozoic Cretaceous Lower middle to late Albian 142 Mesozoic Cretaceous Lower Albian 143 Mesozoic Cretaceous Lower Albian 144 Mesozoic Cretaceous Lower Albian 145 Chénini formation Mesozoic Cretaceous Lower early Albian 146 Chénini formation Mesozoic Cretaceous Lower early Albian

34

Table 2. Dataset containing specimen analyzed in this paper. All data was collected and entered into the dataset as found in the published material it was collected from. If a piece of information was unspecified in the litterature which it came from then information was assumed when possible, this was marked with an asterisk (*). Every row number represents one specimen each.

Row nr. Source Page Image.Label Specimen Dental.Unit 1 Siverson 1997 462 Figure 4A WAM 96.2.42 upper 2 Siverson 1997 462 Figure 4H WAM 96.2.44 lower? 3 Cappetta 2012 255 Figure 233A UM KOB 3 lower* 4 Cappetta 2012 255 Figure 233B UM KOB 4 lower* 5 Cappetta 2012 255 Figure 233E UM KOB 5 lower 6 Cappetta 2012 255 Figure 233F UM KOB 6 upper 7 Benton et al. 2000 242 Fig. 13B lower* 8 Bouaziz et al. 1988 337 Fig. 2K2 lower* 9 Siverson 1997 462 Figure 4O WAM 96.2.75 lower 10 Siverson et al. 2013 92 Fig. 4D NHMUK PV P73050 upper 11 Siverson et al. 2013 94 Fig. 5A NHMUK PV P73059 upper 12 Siverson et al. 2013 94 Fig. 5E NHMUK PV P73060 upper 13 Siverson et al. 2013 94 Fig. 5G NHMUK PV P73061 upper 14 Siverson et al. 2013 94 Fig. 5J NHMUK PV P73062 upper 15 Siverson et al. 2013 94 Fig. 5O NHMUK PV P73063 upper 16 Siverson et al. 2013 94 Fig. 5Q NHMUK PV P73064 upper 17 Siverson et al. 2013 94 Fig. 5U NHMUK PV P73065 upper 18 Siverson et al. 2013 94 Fig. 5Y NHMUK PV P73066 upper 19 Siverson et al. 2013 94 Fig. 5C' NHMUK PV P73067 upper 20 Siverson et al. 2013 94 Fig. 5F' NHMUK PV P73068 upper 21 Siverson et al. 2013 92 Fig. 4H NHMUK PV P73051 upper 22 Siverson et al. 2013 95 Fig. 6A NHMUK PV P73069 lower 23 Siverson et al. 2013 95 Fig. 6O NHMUK PV P73072 lower 24 Siverson et al. 2013 96 Fig. 7H NHMUK PV P73074 lower 25 Siverson et al. 2013 96 Fig. 7M NHMUK PV P73076 lower 26 Siverson et al. 2013 96 Fig. 7T NHMUK PV P73077 lower 27 Siverson et al. 2013 96 Fig. 7W NHMUK PV P73078 lower 28 Siverson et al. 2013 92 Fig. 4L NHMUK PV P73052 upper 29 Siverson et al. 2013 96 Fig. 7C' NHMUK PV P73080 lower 30 Siverson et al. 2013 96 Fig. 7G' NHMUK PV P73081 lower 31 Siverson et al. 2013 98 Fig. 9 (tooth LP3) NHMUK PV P73082 upper* 32 Siverson et al. 2013 98 Fig. 9 (tooth LP9) NHMUK PV P73083 upper* 33 Siverson et al. 2013 97 Fig. 8D SMU 76857 upper* 34 Siverson et al. 2013 97 Fig. 8H SMU 76859 upper* 35 Siverson et al. 2013 97 Fig. 8L SMU 76860 lower* 36 Siverson et al. 2013 92 Fig. 4M NHMUK PV P73053 upper 37 Siverson et al. 2013 92 Fig. 4S NHMUK PV P73054 upper 38 Siverson 1997 462 Figure 4L WAM 96.2.74 upper? 39 Cappetta 2012 257 Figure 235A UM STO 1 lower* 40 Cappetta 2012 257 Figure 235C UM STO 2 lower*

35

41 Cappetta 2012 257 Figure 235F UM STO 3 lower* 42 Cappetta 2012 257 Figure 235I UM KOB 31 lower* 43 Cappetta 2012 257 Figure 235K UM KOB 32 lower* 44 Kriwet et al. 2008 283 Figure 3C MPZ 2005-13 lower* 45 Kriwet et al. 2008 282 Figure 2W MPZ 2005-10 lower* 46 Siverson 1997 462 Figure 4J WAM 96.2.66 lower* 47 Schmitz et al. 2010 286 Fig. 3.1 NLH 102.973 lower* 48 Kriwet 2006 540 Figure 2A BMNH P.36287 lower 49 Cappetta 2012 261 Figure 240D UM PER 2 lower* 50 Schmitz et al. 2010 286 Fig. 3.6 NLH 102.976 lower* 51 Schmitz et al. 2010 286 Fig. 3.9 NLH 102.977 lower* 52 Schmitz et al. 2010 286 Fig. 3.11 NLH 102.978 lower* 53 Schmitz et al. 2010 286 Fig. 3.13 NLH 102.979 upper (?) 54 Cappetta 2012 240 Figure 220C UM TUI 35 lower 55 Antunes and Cappetta 2002 167 Planche 11, Fig. 10 ANG 119 lower 56 Cappetta 2012 232 Figure 214A UM KOB 28 lower* 57 Cappetta 2012 232 Figure 214D UM KOB 29 lower* 58 Cappetta 2012 232 Figure 214F UM KOB 30 lower* 59 Cappetta 2012 241 Figure 221A UM KOB 14 lower* 60 Cappetta 2012 241 Figure 221D UM KOB 15 lower* 61 Siverson 2007 et al. 944 Text-fig. 4A SMU 76311 lower* 62 Siverson 2007 et al. 947 Plate 1, figure 1 SMU 76313 lower* 63 Siverson 2007 et al. 947 Plate 1, figure 20 SMU 76321 lower* 64 Siverson 2007 et al. 947 Plate 1, figure 21 SMU 76322 lower* 65 Siverson 2007 et al. 947 Plate 1, figure 26 SMU 76324 lower* 66 Siverson 2007 et al. 947 Plate 1, figure 4 SMU 76314 lower* 67 Siverson 2007 et al. 947 Plate 1, figure 5 SMU 76315 lower* 68 Siverson 2007 et al. 947 Plate 1, figure 8 SMU 76312 lower* 69 Siverson 2007 et al. 947 Plate 1, figure 10 SMU 76316 lower* 70 Siverson 2007 et al. 947 Plate 1, figure 12 SMU 76317 lower* 71 Siverson 2007 et al. 947 Plate 1, figure 13 SMU 76318 lower* 72 Siverson 2007 et al. 947 Plate 1, figure 16 SMU 76319 lower* 73 Siverson 2007 et al. 947 Plate 1, figure 17 SMU 76320 lower* 74 Siverson 1997 462 Figure 4X WAM 96.2.85 lower* 75 Smart 2007 377 Figure 2A NHM P 66356 lower 76 Smart 2007 377 Figure 2C NHM P 66357 lower 77 Smart 2007 378 Figure 3E NHM P 66361 upper 78 Siverson 2007 et al. 943 Text-fig. 3A SMU 76283 lower* 79 Siverson 2007 et al. 943 Text-fig. 3C SMU 76282 lower* 80 Siverson 2007 et al. 943 Text-fig. 3F SMU 76284 lower* 81 Siverson 2007 et al. 943 Text-fig. 3G SMU 76285 lower* 82 Siverson 2007 et al. 943 Text-fig. 3J SMU 76286 lower* 83 Siverson 2007 et al. 943 Text-fig. 3L SMU 76287 lower* 84 Siverson 2007 et al. 943 Text-fig. 3N SMU 76288 lower* 85 Siverson 2007 et al. 943 Text-fig. 3P SMU 76289 lower*

(continued) 36

Row nr. Side (L=Left, R=Right side of Relative.Position View Order jaw) 1 R anterior labial Lamniformes 2 R? anterior labial Lamniformes 3 parasymphyseal labial Lamniformes 4 anterior labial Lamniformes 5 anterolateral labial Lamniformes 6 labial Lamniformes 7 labial Lamniformes* 8 labial Lamniformes 9 L lateral labial Lamniformes 10 R anterior labial Lamniformes 11 R lateroposterior labial Lamniformes 12 lateroposterior labial Lamniformes 13 L lateroposterior labial Lamniformes 14 L lateroposterior labial Lamniformes 15 L lateroposterior labial Lamniformes 16 L lateroposterior labial Lamniformes 17 L lateroposterior labial Lamniformes 18 R lateroposterior labial Lamniformes 19 R lateroposterior labial Lamniformes 20 R lateroposterior labial Lamniformes 21 L parasymphyseal labial Lamniformes 22 L anterior labial Lamniformes 23 R anterior labial Lamniformes 24 R lateroposterior labial Lamniformes 25 L lateroposterior labial Lamniformes 26 L lateroposterior labial Lamniformes 27 L lateroposterior labial Lamniformes 28 R? anterior labial Lamniformes 29 L lateroposterior labial Lamniformes 30 L lateroposterior labial Lamniformes 31 lateroposterior labial Lamniformes 32 lateroposterior labial Lamniformes 33 L lateroposterior labial Lamniformes 34 L lateroposterior labial Lamniformes 35 L lateroposterior labial Lamniformes 36 parasymphyseal labial Lamniformes 37 R anterior labial Lamniformes 38 R? lateral labial Lamniformes 39 anterior labial Lamniformes 40 anterolateral labial Lamniformes 41 anterolateral labial Lamniformes 42 anterior labial Lamniformes 43 anterolateral labial Lamniformes 44 lateral lingual Lamniformes 45 posterior labial Lamniformes

37

46 labial Lamniformes 47 anterior labial Lamniformes 48 lateral lingual Lamniformes 49 anterolateral labial Lamniformes 50 anterior labial Lamniformes 51 lateral labial Lamniformes 52 posterolateral labial Lamniformes 53 parasymphyseal labial Lamniformes 54 anterior labial Lamniformes 55 anteriolateral labial Lamniformes 56 anterior labial Lamniformes 57 lateral labial Lamniformes 58 anterolateral labial Lamniformes 59 anterior labial Lamniformes 60 anterolateral labial Lamniformes 61 lateral labial Lamniformes 62 anterior labial Lamniformes 63 lateroposterior labial Lamniformes 64 lateroposterior labial Lamniformes 65 posterior labial Lamniformes 66 anterolateral labial Lamniformes 67 lateral labial Lamniformes 68 lateral labial Lamniformes 69 lateroposterior labial Lamniformes 70 lateral labial Lamniformes 71 lateral labial Lamniformes 72 lateroposterior labial Lamniformes 73 lateral labial Lamniformes 74 lateral labial Lamniformes 75 L lateral labial Lamniformes 76 R lateral labial Lamniformes 77 L lateral labial Lamniformes 78 anterolateral labial Lamniformes 79 lateral labial Lamniformes 80 lateral labial Lamniformes 81 lateral labial Lamniformes 82 lateral labial Lamniformes 83 lateroposterior labial Lamniformes 84 lateral labial Lamniformes 85 posterior labial Lamniformes

(continued) Row nr. Family Genus Species Continent Country 1 Cretoxyrhinidae Archaeolamna sp. Australia Australia 2 Cretoxyrhinidae Archaeolamna sp. Australia Australia 3 Cretodus semiplicatus Asia Kazakhstan (central)

38

4 Cretodus semiplicatus Asia Kazakhstan (central) 5 Cretodus semiplicatus Asia Kazakhstan (central) 6 Cretodus semiplicatus Asia Kazakhstan (central) 7 Cretodus Africa Tunisia 8 Cretoxyrhinidae Cretodus ? Africa Tunisia 9 Cretoxyrhinidae Cretolamna sp. Australia Australia 10 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 11 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 12 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 13 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 14 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 15 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 16 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 17 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 18 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 19 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 20 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 21 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 22 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 23 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 24 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 25 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 26 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 27 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 28 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 29 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 30 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 31 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 32 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 33 Cretoxyrhinidae Cretoxyrhina vraconensis North USA America 34 Cretoxyrhinidae Cretoxyrhina vraconensis North USA America 35 Cretoxyrhinidae Cretoxyrhina vraconensis North USA America 36 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 37 Cretoxyrhinidae Cretoxyrhina vraconensis Asia Kazakhstan (central) 38 Cretoxyrhinidae incertae sedis Australia Australia 39 Dwardius siversoni Europe Russia 40 Dwardius siversoni Europe Russia 41 Dwardius siversoni Europe Russia 42 Dwardius sp. Asia Kazakhstan (central)

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43 Dwardius sp. Asia Kazakhstan (central) 44 Eoptolamnidae Eoptolamna eccentrolopha Europe Spain 45 Eoptolamnidae Eoptolamna eccentrolopha Europe Spain 46 Cretoxyrhinidae Leptostyrax sp. Austalia Australia 47 Eoptolamnidae Leptostyrax stychi sp. nov. Europe Germany 48 incertae sedis Priscusurus adruptodontus sp. South Peru nov. America 49 Priscusurus adruptodontus South Peru America 50 Eoptolamnidae Protolamna sarstedtensis sp. Europe Germany nov. 51 Eoptolamnidae Protolamna sarstedtensis sp. Europe Germany nov. 52 Eoptolamnidae Protolamna sarstedtensis sp. Europe Germany nov. 53 Eoptolamnidae Protolamna sarstedtensis sp. Europe Germany nov. 54 Pseudoscapanorhynchidae Protolamna sokolovi Europe France 55 Cretoxyrhinidae Protolamna sp. Africa Angola 56 Cardabiodontidae Pseudoisurus aff. tomosus Asia Kazakhstan (central) 57 Cardabiodontidae Pseudoisurus aff. tomosus Asia Kazakhstan (central) 58 Cardabiodontidae Pseudoisurus aff. tomosus Asia Kazakhstan (central) 59 Pseudoscapanorhynchidae Pseudoscarpanorhynchus aff. compressidens Asia Kazakhstan (central) 60 Pseudoscapanorhynchidae Pseudoscarpanorhynchus aff. compressidens Asia Kazakhstan (central) 61 Anacoracidae Squalicorax aff. S. baharijensis North USA America 62 Anacoracidae Squalicorax pawpawensis sp. North USA nov. America 63 Anacoracidae Squalicorax pawpawensis sp. North USA nov. America 64 Anacoracidae Squalicorax pawpawensis sp. North USA nov. America 65 Anacoracidae Squalicorax pawpawensis sp. North USA nov. America 66 Anacoracidae Squalicorax pawpawensis sp. North USA nov. America 67 Anacoracidae Squalicorax pawpawensis sp. North USA nov. America 68 Anacoracidae Squalicorax pawpawensis sp. North USA nov. America 69 Anacoracidae Squalicorax pawpawensis sp. North USA nov. America 70 Anacoracidae Squalicorax pawpawensis sp. North USA nov. America 71 Anacoracidae Squalicorax pawpawensis sp. North USA nov. America 72 Anacoracidae Squalicorax pawpawensis sp. North USA nov. America 73 Anacoracidae Squalicorax pawpawensis sp. North USA nov. America 74 Anacoracidae Squalicorax primaevus Australia Australia 75 Anacoracidae Squalicorax primaevus Europe England 76 Anacoracidae Squalicorax primaevus Europe England 77 Anacoracidae Squalicorax primaevus Europe England 78 Anacoracidae Squalicorax priscoserratus sp. North USA nov. America 79 Anacoracidae Squalicorax priscoserratus sp. North USA nov. America 80 Anacoracidae Squalicorax priscoserratus sp. North USA nov. America 81 Anacoracidae Squalicorax priscoserratus sp. North USA nov. America 82 Anacoracidae Squalicorax priscoserratus sp. North USA

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nov. America 83 Anacoracidae Squalicorax priscoserratus sp. North USA nov. America 84 Anacoracidae Squalicorax priscoserratus sp. North USA nov. America 85 Anacoracidae Squalicorax priscoserratus sp. North USA nov. America

(continued) Row nr. Formation Era Period Epoch Sub Age 1 Mesozoic Cretaceous Lower middle to late Albian 2 Mesozoic Cretaceous Lower middle to late Albian 3 Mesozoic Cretaceous Lower late Albian 4 Mesozoic Cretaceous Lower late Albian 5 Mesozoic Cretaceous Lower late Albian 6 Mesozoic Cretaceous Lower late Albian 7 Chenini formation Mesozoic Cretaceous Lower early Albian 8 Chénini formation Mesozoic Cretaceous Lower early Albian 9 Mesozoic Cretaceous Lower middle to late Albian 10 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 11 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 12 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 13 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 14 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 15 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 16 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 17 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 18 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 19 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 20 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 21 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 22 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 23 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 24 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 25 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 26 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 27 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 28 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 29 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 30 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 31 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 32 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 33 Pawpaw formation Mesozoic Cretaceous Lower late Albian 34 Pawpaw formation Mesozoic Cretaceous Lower late Albian

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35 Pawpaw formation Mesozoic Cretaceous Lower late Albian 36 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 37 Mesozoic Cretaceous Lower latest late/earliest Albian/Cenomanian early 38 Mesozoic Cretaceous Lower middle to late Albian 39 Mesozoic Cretaceous Lower Albian 40 Mesozoic Cretaceous Lower Albian 41 Mesozoic Cretaceous Lower Albian 42 Mesozoic Cretaceous Lower late Albian 43 Mesozoic Cretaceous Lower late Albian 44 Artoles formation Mesozoic Cretaceous Lower late Barremian 45 Artoles formation Mesozoic Cretaceous Lower late Barremian 46 Mesozoic Cretaceous Lower middle to late Albian 47 Mesozoic Cretaceous Lower early Barremian 48 Muerto Limestone Mesozoic Cretaceous Lower middle? Albian formation 49 Muerto Limestone Mesozoic Cretaceous Lower middle Albian formation 50 Mesozoic Cretaceous Lower early Barremian 51 Mesozoic Cretaceous Lower early Barremian 52 Mesozoic Cretaceous Lower early Barremian 53 Mesozoic Cretaceous Lower early Barremian 54 Mesozoic Cretaceous Lower middle to late Albian 55 Mesozoic Cretaceous Lower Albian 56 Mesozoic Cretaceous Lower late Albian 57 Mesozoic Cretaceous Lower late Albian 58 Mesozoic Cretaceous Lower late Albian 59 Mesozoic Cretaceous Lower late Albian 60 Mesozoic Cretaceous Lower late Albian 61 Pawpaw formation Mesozoic Cretaceous Lower late Albian 62 Pawpaw formation Mesozoic Cretaceous Lower late Albian 63 Pawpaw formation Mesozoic Cretaceous Lower late Albian 64 Pawpaw formation Mesozoic Cretaceous Lower late Albian 65 Pawpaw formation Mesozoic Cretaceous Lower late Albian 66 Pawpaw formation Mesozoic Cretaceous Lower late Albian 67 Pawpaw formation Mesozoic Cretaceous Lower late Albian 68 Pawpaw formation Mesozoic Cretaceous Lower late Albian 69 Pawpaw formation Mesozoic Cretaceous Lower late Albian 70 Pawpaw formation Mesozoic Cretaceous Lower late Albian 71 Pawpaw formation Mesozoic Cretaceous Lower late Albian 72 Pawpaw formation Mesozoic Cretaceous Lower late Albian 73 Pawpaw formation Mesozoic Cretaceous Lower late Albian 74 Mesozoic Cretaceous Lower middle to late Albian 75 Mesozoic Cretaceous Lower middle Albian 76 Mesozoic Cretaceous Lower middle Albian 77 Mesozoic Cretaceous Lower middle Albian 78 Pawpaw formation Mesozoic Cretaceous Lower late Albian 79 Pawpaw formation Mesozoic Cretaceous Lower late Albian

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80 Pawpaw formation Mesozoic Cretaceous Lower late Albian 81 Pawpaw formation Mesozoic Cretaceous Lower late Albian 82 Pawpaw formation Mesozoic Cretaceous Lower late Albian 83 Pawpaw formation Mesozoic Cretaceous Lower late Albian 84 Pawpaw formation Mesozoic Cretaceous Lower late Albian 85 Pawpaw formation Mesozoic Cretaceous Lower late Albian

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