THE PENNSYLVANIA STATE UNIVERSITY SCHREYER HONORS COLLEGE

DEPARTMENT OF SCIENCE

EARLY CAMPANIAN ELASMOBRANCH ASSEMBLAGE FROM NORTH CAROLINA WITH PALEOECOLOGICAL AND PALEOBIOGEOGRAPHICAL IMPLICATIONS

TAYLOR A. KIGHTLINGER SPRING 2017

A thesis submitted in partial fulfillment of the requirements for a baccalaureate degree in General Biology with honors in Biology

Reviewed and approved* by the following:

Todd Cook Professor of Paleontology Thesis Supervisor

Michael Campbell Professor of Cell Biology Honors Adviser

* Signatures are on file in the Schreyer Honors College. i

ABSTRACT

Recently, a new sclerorhynchid sawfish, Borodinopristis shannoni, was described from an early

Campanian (± 80 million years ago) fossil site near Elizabethtown, Bladen County, North Carolina, USA.

At the time of recovery, a large assemblage of additional elasmobranch remains, mostly teeth, was also collected. Preliminary examination of the fossil material revealed a rich diversity of shark and ray species in this region of the eastern seaboard, pertaining to this particular stage of the Cretaceous period, that had not been previously well-assessed.

By comparing this Elizabethtown assemblage to slightly younger assemblages, changes in regional faunal composition over a relatively short amount of geological time were inferred. These

Elizabethtown fossils were also evaluated with contemporaneous assemblages from Europe, northern

Africa, and Australia to determine whether a species had cosmopolitan or regional ranges.

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TABLE OF CONTENTS

LIST OF FIGURES ...... iii

ACKNOWLEDGEMENTS ...... v

Chapter 1 Introduction ...... 1

Sub-Chapter 1 Geology ...... 2 Sub-Chapter 2 Materials & Methods ...... 3

Chapter 2 Systematics ...... 4

Lonchidion babulski ...... 4 Hybodus sp...... 4 Squantina hassei ...... 4 Plicatoscyllium globidens ...... 5 Scapanorhynchous texanus ...... 5 Cretalamna appendiculata ...... 6 Archeolamna kopingensis ...... 6 Squalicorax kaupi ...... 6 Squalicorax pristidontis ...... 7 Ischyrhiza mira ...... 7 Pseudohypolophus ellipsis ...... 7 Rhombodus laevis ...... 8 Brachyrhizodus wichitaensis ...... 8

Chapter 3 Discussion ...... 12

LITERATURE CITED ...... 14

iii

LIST OF FIGURES

Figure 1. Stratigraphy of Black Creek ...... 2

Figure 2. Elasmobranchs ...... 9

Figure 3. Lamniformes ...... 10

Figure 4. Batoids ...... 11

iv

ACKNOWLEDGEMENTS

I express my utmost gratitude to Gerard Case for providing the fossil specimens, to Jerome

Magraw of Penn State Behrend for his assistance with the SEM imaging, and to Dr. Michael Campbell of

Penn State Behrend, my honors advisor, for reviewing an early draft. The completion of this thesis would not have been possible without Dr. Todd Cook, my research advisor. He has continually offered encouragement toward my success on this project, my education, and overall career goes. I bestow my sincerest appreciation for the learning and publication opportunities that the Penn State Behrend 2016

Undergraduate Student Research Grant has given me. I am grateful for the supportive School of Science staff at Penn State Behrend, as well.

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Chapter 1

Introduction

Chondrichthyans (cartilaginous fishes) include the elasmobranchs (sharks, rays, skates, and sawfish) and holocephalans (chimaeras). The fossil record of this group is well-known but is largely restricted to hard elements that preserve readily, such as teeth, scales, and spines. The dental morphology of elasmobranchs is species specific and is used in identification process and to provide paleoecological and paleobiogeographical insights.

Recently, a new sclerorhynchid sawfish, Borodinopristis shannoni, was recovered from a productive early Campanian (ca. 80 Ma) fossil site near Elizabethtown, Bladen County, North Carolina,

USA (Case et al., 2012). At the time of recovery, a large assemblage of additional elasmobranch remains

(largely teeth) was also collected. Very little is known of shark and ray diversity in this region of the

Eastern seaboard during this particular stage of the Cretaceous period. Cicimurri (2007) described a late

Campanian (ca. 76 Ma) assemblage from South Carolina that included species belonging to 18 different genera. By comparing the latter assemblage with the recovered assemblage from Elizabethtown, we will examine the changes in regional faunal composition over a relatively short amount of geological time. By comparing the occurrence of species at Elizabethtown with contemporaneous assemblages from Europe, northern Africa, and Australia, we will be able to determine whether species had cosmopolitan or regional ranges. 2

Sub-Chapter 1 Geology

The elasmobranch assemblage was recovered from the Bladen Formation in an exposed landfill pit near Elizabethtown, Bladen County, North Carolina, USA. This formation, which overlies the Tar

Heel Formation and underlies the Donoho Creek Formation, represents a delta-shelf system and largely consists of feldspathic quartz sandstone and dark clay (Sohl and Owens 1991). Based on palynological data, the Bladen Formation is likely middle Campanian in age (Prowell et al. 2003).

Figure 1. Geographical and stratigraphic map of the Elizabethtown site (image taken from Case et al., 2012)

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Sub-Chapter 2 Materials & Methods

Teeth and rostral spine specimens were collected by sieving and handpicking. To dissolve remaining sediment or matrices, they were immerse in an acetic acid solution. Considerably small teeth were gold-coated and imaged using a Hitachi S-2500 Scanning Electron Microscope. Larger specimens were coated with ammonium chloride and photographed using a Canon EOS Rebel T6 50mm compact- macro lens camera. 4

Chapter 2

Systematics

Chondrichthyes Huxley, 1880 Elasmobranchii Bonaparte, 1838 Euselachii Hay, 1902 Hybodontiformes Maisey, 1975 Lonchidion Estes, 1964 Lonchidion sp. (Fig. 2; A1, A2)

The crown mesodistally elongated and labiolingually flat. Both labial and lingual crown faces are smooth. There is a moderate central protuberance on the labial crown face. The root is missing.

Hybontidae Owen, 1846 Hybodus Agassiz, 1837 Hybodus sp. (Fig. 2; B1, B2)

The crown consists of a tall median cusp and a two pairs of small lateral cusplets. Both labial and lingual faces are convex and separated by a smooth cutting edge. Strong enameloid folds extend from the base to approximately half the median cusp height. Folds associated with the lateral cusplets extend to the apex.

The root is missing.

Neoselachii Compagno, 1977 Squalomorphii Compagno, 1973 Squantiformes Buen, 1926 Squantinidae Bonaparte, 1838 Squantina Dumeril, 1806 Squantina hassei Leriche, 1929 5 (Fig. 2; C1, C2)

The crown contains a triangular median cusp with a sharp apex and a wide base. The labial and lingual crown faces are smooth and separated by a cutting edge. The heels are oblique and separated from the cusp. The base of the labial crown has a well-developed apron. The lingual crown face has a lingual protuberance. The root is damaged but is relatively triangular basally and has a large central foramen.

Orectolobiformes Applgegate, 1972 Ginglymostomatidae Gill, 1862 Plicatoscyllium Case and Cappetta, 1997 Plicatoscyllium globidens Capetta and Case, 1975 (Fig. 2; D1, D2)

The crown contains a lingual directed median cusp and one pair of short lateral cusplets. The labial face contains enameloid folding and has a broad apron basally. The lingual face is smooth and has a medio- lingual protuberance. The root has elongated root lobes that are V-shaped and a large central foramen.

Lamniformes Berg, 1937 Mitsukurinidae Jordan, 1898 Scapanorhynchus Woodward, 1889 Scapanorhynchus texanus Roemer, 1849 (Fig. 3; A1, A2, B1, B2)

The anterior tooth crown contains a tall median cusp with smooth cutting edges. The labial crown is flat and more or less smooth. The lingual crown is convex and contains enameloid folds that extend from the base to over half the height of the cusp. A sharp cutting edge is present. There is a prominent lingual protuberance that has a well-developed nutrient groove that possesses three foramina. The root contains two elongated root lobes that are separated by a deep basal concavity.

The crown of the lateral tooth has tall distally inclined median cusp and two pairs small of lateral cusplets. The cutting edge is smooth. The smooth labial crown face is flat, whereas the smooth lingual crown face is convex. The lingual protuberance is also divided by a nutrient groove that contains central 6 foramina. The root contains two wide root lobes that are separated by a narrow basal concavity.

Otodontidae Glikman, 1964 Cretalamna Glikman, 1958 Cretalamna appendiculata Agassiz, 1843 (Fig. 3; C1, C2)

The crown contains a broad distally inclined median cusp and pair of broad lateral cusplets. The labial crown face is flat, whereas the lingual face is convex. Both faces are smooth and separated by a smooth cutting edge. The root is somewhat eroded and has a weakly developed lingual protuberance. The root lobes are relatively broad and separated by a shallow basal concavity.

Archaeolamnidae Underwood and Cumbaa, 2010 Archaeolamna Siverson, 1992 Archaeolamna kopingensis Siverson, 1992 (Fig. 3; D1, D2)

The crown contains a tall median cusp and one pair of narrow lateral cusplets. The cutting edges are smooth. The labial crown face is more or less flat but has a distinct indentation extending from the base to approximately one-third the height of the median cusp. The lingual crown face is convex and smooth. The root is somewhat eroded but has a lingual protuberance. The elongated root lobes are separated by a deep basal concavity. There is a central foramen positioned in the root underneath the indentation. The lingual crown contains few striations and has a large protuberance. The root includes two concaved lobes.

Anacoracidae Casier, 1947 Squalicorax Whitley, 1939 Squalicorax kaupi Agassiz, 1843 (Fig. 3; E1, E2)

The crown contains a median cusp that is broad and distally inclined. The labial crown face is flat, whereas the lingual crown face is convex. Both faces are smooth. The mesial cutting edge in strongly convex and contains distinct serrations that decrease in size towards the apex and base. The distal cutting 7 edge is vertically directed, straight and contains relatively similar sized serrations. The distal heel is obliquely directed and contains serrations that decrease in size basally. The root contains a large protuberance and root lobes separated by a deep basal concavity. The root is porous containing numerous foramina.

Squalicorax pristodontus Agassiz, 1843 (Fig. 3; F1, F2)

The crown of this tooth has a notched distal edge. The medial edge extends diagonally and then curves laterally until it reaches the root. The cutting edges are slightly serrated into equal portions. The labial crown is contains deep striations and few ridges. The lingual crown is smaller in size, due to erosion. The labial root has foramina just below the crown that extend laterally and fully across the tooth.

Sclerorhynchiformes Kriwet, 2004 Sclerorhynchidae Cappetta, 1974 Ischyrhiza Leidy, 1856 Ischyrhiza mira Leidy, 1856 (Fig. 4; A1, A2)

The rostral spine is contains convex dorsal and ventral surfaces that are separated by distinct anterior and posterior cutting edges. The spine is slightly ventrally inclined. The peduncle large and expands basally forming lobes that are separated by a concavity.

Rajiformes Berg, 1937 Rhinobatoidei incertae familiae Pseudohypolophus Cappetta and Case, 1975 Pseudohypolophus ellipsis Case, Schwimmer, Borodin and Leggett, 2001 (Fig. 4; B1, B2)

The bulbous hexagonal crown is tall and smooth and overhangs the root. The occlusal, labial, and lingual faces are slightly convex, whereas the margino-labial and margino-lingual faces are 8 slightly concave. The root has numerous foramina located below the crown. A distinct nutrient groove separates the root lobes and also contains foramina.

Myliobatiformes Campagno, 1973 Rhombodontidae Cappetta, 1987 Rhombodus Dames, 1881 Rhombodus laevis Cappetta and Case, 1975 (Fig. 4; C1, C2)

The rhomboid crown is tall and smooth. The occlusal face is flat, whereas the margino-labial and margino-lingual faces are very slightly concave. The root is tall and contains numerous foramina. The root lobes are separated from each other by a distinct nutrient groove.

Myliobatidae Bonaparte, 1838 Brachyrhizodus Romer, 1942 Brachyrhizodus wichitaensis Romer, 1942 (Fig. 4; D1, D2)

The hexagonal crown is tall and mesodistally elongated. The occusal, labial, and lingual faces are flat and smooth. The root is contains five root loblets that are separated from each other by distinct grooves.

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Figure 2. Hybodontiform, squatiniform, and orectolobiform species recovered from the

Elizabethtown site. Scale bar represents 1mm.

10

Figure 3. Lamniform species recovered from the Elizabethtown site. Scale bar represents

1mm.

11

Figure 4. Batoid species recovered from the Elizabethtown site. Scale bar represents 1mm. 12

Chapter 3

Discussion

This shallow water marine environment assemblage includes 13 species from 12 genera, at least

11 families, and 6 orders. Extant lamniforms (mackerel sharks) are large apex predators with large biogeographical ranges (Musick et al. 2004). The lamniform species recovered from this assemblage also had very wide paleobiogeographical distributions. Scapanorhynchous texanus has been recovered from

Campanian and Maastrichtian deposits of North America (Welton and Farish 1993; Cappetta and Case

1999; Becker et al. 2008; Bourdon et al. 2011), the Near East (Mustafa et al. 2002) and Africa (Antunes and Cappetta 2002). Cretalamna appendiculata has been reported from Albian to upper Eocene of North

America, South America, Europe, the Near East, and northern Africa (Cappetta 2012). Archaeolamna kopingensis has been recovered Albian to Maastricthian deposits of Europe (Davis 1890; Siverson 1992;

Biddle 1993; Vullo 2007), Australia (Siverson, 1996; 1997) and the Atlantic and Gulf coastal plains and

Western Interior Seaway of North America (Cappetta and Case 1975; Beavan and Russell 1999; Becker et al. 2000; Cumbaa et al. 2006; Cook et al. 2008, 2011, 2013; Shimada and Martin 2008; Underwood and

Cumbaa 2010). Both Squalicorax kaupi and S. pristodontus are known from Coniacian to Maastrichtian deposits of Europe (Cappetta 2012; Sørensen et al. 2013), the Near East (Zalmont and Mustafa 2001),

Africa (Gottfried et al. 2001), South America (Da Silva 2007), and the Atlantic and Gulf coastal plains and Western Interior Seaway of North America (Cappetta and Case 1975; Welton and Farish 1993; Case and Cappetta 1997; Cappetta and Case 1999; Case et al. 2001; Becker et al. 2006; Shimada and Cicimurri

2006; Cuny et al. 2012; Corral et al. 2016). The orectolobiform (carpet sharks) Plicatoscyllium globidens has been recovered from Campanian and Maastrichtian deposits from the Atlantic and Gulf coastal plains

(Cappetta and Case 1975; Case 1979; Case and Schwimmer 1988; Case 1991) and the Western Interior

Seaway (Case 1987) and has only been reported from France (Vullo 2005). Extant squatiniforms (angel 13 sharks) are benthic and have limited vagility (Musick 2004), therefore the presence of Squatina hassei from Campanian and Maasstrichtian deposits of North America (Capetta and Case 1975; Welton and

Farish 1993; Case et al. 2001; Schwimmer et al. 2001) and Europe (Vullo 2005) is surprising. The myliobatiform (stingray) species Brachyrhizodus wichitaensis has been recovered from Campanian and

Maastrichtian deposits of North America (Cappetta and Case 1975; Welton and Farish 1993) and Africa

(Antunes and Cappetta 2002).

Many of the species of this assemblage were restricted to the North American marine waters. The sclerorhynchiform (extinct sawfish) Ischyrhiza mira, the rajiform (extinct guitarfish) Pseudohypolophus ellipsis, and the myliobatiform Rhombodus laevis and the hybodont Lonchidion babulskii has been reported from Turonian to Maastrichtian deposits of the Atlantic and Gulf coastal plains (Case 1991; Case

1995; Becker et al. 2006) and the Western Interior Seaway (Beavan and Russell 1999; Bourdon et al.

2011). The species with the most restricted distribution is Hybodus novojerseyensis that has only been previously recovered from New Jersey (Case and Cappetta 2004).

In comparison to species from a slightly younger assemblage from the Donoho Creek

Formation of South Carolina (Cicimurri, 2007), the change in fauna from middle Campanian to late Campanian was miniscule. Euselachian species shared by both assemblages include

Hybodus sp., Lonchidion babulskii, Squalicorax kaupi, Squalicorax pristodontus, Archaeolamna kopingensis, Cretalamna appendiculata, Rhombodus laevis, Brachyrhizodus wichitaensis, and

Ischyrhiza mira. Some taxa, such as Scapanorhynchous texanus, are absent from the Donho Creek assemblage. However, the Donoho Creek Formation contains not only marine fossils, but also terrestrial specimens due to its prodelta deposition. 14

LITERATURE CITED

Agassiz, J.L.R. 1833–1844. Recherches sur les poissons fossils, 3. Imprimerie de Petitpierre,

Neuchâtel.

Antunes, M.T. and H. Cappetta. 2002. Sélaciens du Crétacé (Albien-Maastrichtien) d'Angola.

Palaeontographica, Abt. A 264:85–146.

Applegate, S.P. 1972. A revision of the higher taxa of orectolobids. Journal of the Marine

Biology Association of India, 14:743–751.

Beavan, N.R. and A.P. Russell. 1999. An elasmobranch assemblage from the terrestrial-marine

transitional Lethbridge Coal Zone (Dinosaur Park Formation: Upper Campanian),

Alberta, Canada. Journal of Paleontology, 73:494–503.

Becker, M.A., J.A. Chamberlain, and P.W. Stoffer. 2000. Pathologic tooth deformities in

modern and fossil chondrichthians: a consequence of feeding-related injury. Lethaia,

33:103–118.

Becker, M.A., J.A. Chamberlain, and D.O. Terry. 2004. Chondrichthyans from the Fairpoint

Member of the Fox Hills Formation (Maastrichtian), Meade County, South Dakota.

Journal of Vertebrate Paleontology, 24:780–793.

Becker, M.A., J.A. Chamberlain, and G.E. Wolf. 2006. Chondrichthyans from the Arkadelphia

Formation (Upper Cretaceous: upper Maastrichtian) of Hot Spring County, Arkansas.

Journal of Paleontology, 80:700–716. 15 Becker, M.A., R.B. Chamberlain, and J.A. Chamberlain. 2008. Large carcharhinoid-type shark

vertebrae in the Upper Cretaceous of New Jersey: evidence for an anacoracid origin.

Northeastern Geology and Environmental Science, 30:118–129.

Berg, L.S. 1937. A classification of fish-like vertebrates. Bulletin de l’Académie des Sciences de

l'URSS. Classe des Sciences Mathématiques et Naturelles, 1937:1277–1280.

Biddle, J.P. 1993. Les Elasmobranches de l´Albien inférieur et moyen (Crétacé inférieur) de la

Marne et de la Haute-Marne (France). Belgian Geological Survey, Professional Paper:

Elasmobranches et Stratigraphie, 264:191–240.

Bonaparte, C.L. 1838. Selachorum tabula analytica. Nuovi Annali della Scienze Naturali,

Bologna, 1:195–214.

Buen, F. 1926. Catalogo ictiologico del Mediterraneo Español y de Marruecos, recopilando lo

publicado sobrepeces de las costas mediterraneas y proximas del Atlantico (Mar de

España). Resultados de las ampafias Realizadas por Acuerdos Internacionales. Instituto

Español de Oceanografia, 2:1–221.

Cappetta, H. 1974. Sclerorhynchidae nov. fam., Pristidae et Pristiophoridae: un exemple de

parallélisme chez les Sélaciens. Comptes Rendus hebdomadaires des séances de

l'Academie des Sciences, 278:225–228.

Cappetta, H. 1987. Handbook of Paleoichthyology, Vol. 3B: Chondrichthyes II. Gustav Fischer

Verlag, Stuttgart.

Cappetta, H. 2012. Chondrichthyes. Mesozoic and Cenozoic Elasmobranchii: Teeth. Handbook

of Palaeoichthyology, Volume 3E. München: Verlag Dr. Friedrich Pfeil.

Cappetta, H. and G.R. Case. 1975. Contribution à l'étude des Sélaciens du groupe Monmouth

(Campanien-Maestrichtien) du New Jersey. Palaeontographica, Abt. A, 151:1–46. 16 Cappetta, H. and G.R. Case. 1999. Additions aux faunes de sélaciens du Crétacé du Texas

(Albien supérieur-Campanien). Palaeo Ichthyologica, 9:5–111.

Case, G.R. 1979. Cretaceous selachians from the Peedee Formation (Late Maastrichtian) of

Duplin Country, North Carolina. Brimleyana, 2:77–89.

Case, G.R. 1987. A new selachian fauna from the Late Campanian of Wyoming (Teapot

Sandstone Member, Mesaverde Formation, Big Horn Basin). Palaeontographica, Abt. A,

197:1–37.

Case, G.R. 1995. Fossil shark remains from the early and middle Maastrichtian of the Upper

Cretaceous of Monmouth County, New Jersey. Pp. 72–80 in J. Baker (ed). Contributions

to the Paleontology of New Jersey. Geological Association of New Jersey.

Case, G.R. and D.R. Schwimmer. 1988. Late Cretaceous fish from the Blufftown Formation

(Campanian) in western Georgia. Journal of Paleontology, 62:290–301.

Case, G.R. and H. Cappetta. 1997. A new selachian fauna from the Late Maastrichtian of Texas

(Upper Cretaceous/Navarroan; Kemp Formation). Müenchner Geowissenschaftliche

Abhandlungen Reihe A Geologie und Paläeontologie, 34:131–189.

Case, G.R. and H. Cappetta. 2004. Additions to the elasmobranch fauna from the late Cretaceous

of New Jersey (lower Navesink Formation, early Maastrichtian). Palaeovertebrata, 33:1–

16.

Case, G.R., P.D. Borodin and J.J. Leggett. 2001. Fossil selachians from the New Egypt

Formation (Upper Cretaceous, Late Maastrichtian) of Arneytown, Monmouth County,

New Jersey. Palaeontographica, Abt. A, 261:113–124. 17 Casier, E. 1947. Constitution et évolution de la racine dentaire des Euselachii. II. Etude

comparative des types. Bulletin du Musée Royal d'Histoire Naturelle de Belgique, 23:1–

32.

Compagno, L.J.V. 1973. Interrelationships of living elasmobranchs. Pp. 15–61 in P.H.

Greenwood, R.S. Miles and C. Patterson. Interrelationships of Fishes. Zoological Journal

of the Linnean Society 53.

Compagno, L.J.V. 1977. Phyletic relationships of living sharks and rays. American Zoologist,

17:303−322.

Cook, T.D., M.V.H. Wilson, and A.M. Murray. 2008. A middle Cenomanian euselachian

assemblage from the Dunvegan Formation of northwestern Alberta. Canadian Journal of

Earth Sciences, 45:1185–1197.

Cook, T.D., M.G. Newbrey, A.M. Murray, M.V.H. Wilson, K. Shimada, G.T. Takeuchi, and

J.D. Stewart. 2011. A Partial Skeleton of the Late Cretaceous Lamniform Shark,

Archaeolamna kopingensis, from the Pierre Shale of Western Kansas, U.S.A. Journal of

Vertebrate Paleontology, 31:8–21.

Cook, T.D., M.V.H Wilson, A.M. Murray, A.G. Plint, M.G. Newbrey, and M.J. 2013. A high

latitude euselachian assemblage from the early Turonian of Alberta, Canada. Journal of

Systematic Palaeontology, 11:555–587.

Corral, J.C., A. Berreteaga, and H. Cappetta. 2016. Upper Maastrichtian shallow marine

environments and neoselachian assemblages in North Iberian palaeomargin (Castilian

Ramp, Spain). Cretaceous Research, 57:639–661. 18 Cumbaa, S.L., C. Schröder-Adams, R.G. Day, and A.J. Phillips. 2006. Cenomanian bonebed

faunas from the northeastern margin, Western Interior seaway, Canada. New Mexico

Museum of Natural History and Science, Bulletin, 35:139–155.

Cuny, G., J.E. Martin and R. Sarr. 2012. A neoselachian shark fauna from the Late Cretaceous

of Senegal. Cretaceous Research, 34:107–115.

Dames, W. 1881. Über Fischzähne aus der obersenonen Tuffkreide von Maastricht für welcher

den Gattungsnamen Rhombodus vorschlug. Sitzungsberichte der Gesellschaft

naturforschender Freunde zu Berlin 1881:1–3.

Da Silva, C. and M. Bürgener. 2007. South Africa's demersal shark meat harvest. Traffic

Bulletin, 21:55–66.

Davis, J.W. 1890. On the fossil fish of the Cretaceous formations of Scandinavia. Scientific

Transactions of the Royal Dublin Society, 2:363–434.

Dumeril, A.H.A. 1806. Zoologie analytique, ou méthode naturelle de classification des animaux.

Paris: 1–344.

Estes, R. 1964. Fossil vertebrates from the Late Cretaceous Lance Formation, eastern Wyoming.

University of California Publications in Geological Sciences, 49:1–180.

Gill, T.N. 1862. Analytical synopsis of the Order of Squali and revision of the nomenclature of

the genera. Annals of the Lyceum of Natural History of New York, 7:367–408.

Glikman, L.S. 1958. Rates of evolution in Lamnoid sharks. Doklady Akademij Nauk SSSR

123:568–571. [In Russian].

Glikman, L.S. 1964. Sharks of Paleogene and their stratigraphic significance. Nauka Press. [In

Russian]. 19 Gottfried, M.D., J.A. Rabarison and L.L. Randriamiarimanana. 2001. Late Cretaceous

elasmobranchs from the Mahajanga Basin of Madagascar. Cretaceous Research, 22:491–

496.

Hay, O.P. 1902. Bibliography and catalogue of the fossil Vertebrata in North America. United

States Geological Survey Bulletin, 179:1–868.

Huxley, T.H. 1880. On the application of the laws of evolution to the arrangement of the

Vertebrata and more particularly of the Mammalia. Proceedings of the Zoological Society

of London, 1880:649–662.

Jordan, D.S. 1898. Description of a species of fish (Mitsukurina owstoni) from Japan, the type of

a distinct family of Lamnoid sharks. Proceedings of the California Academy of Sciences,

1:199–202.

Kriwet J. 2004. The systematic position of the Cretaceous sclerorhynchid sawfishes

(Elasmobranchii, Pristiorajea). Mesozoic Fishes 3 – Systematics, Paleoenvironments and

Biodiversity:57–73.

Kriwet, J. 2004. Late Jurassic selachians (Chondrichthyes: Hybodontiformes, Neoselachii) from

Central Portugal. Neues Jahrbuch für Geologie und Palaontologie, Monatshefte,

2004:233–256.

Leidy, J. 1856. Notice of remains of extinct vertebrate animals of New Jersey, collected by Prof.

Cook of the State Geological Survey under the direction of Dr. W. Kitchel. Proceedings

of the Academy of Natural Sciences of Philadelphia, 8:220–221.

Leriche, M. 1929. Sur une forme nouvelle du genre Chlamydoselachus (C. tobleri) rejetée par le

volcan de boue de Chagonary (île de la Trinité, Petites-Antilles). Bulletin de la Société

Belge de Géologié, de Paléontologie et d'Hydrologie, 38:55–58. 20 Maisey, J.G. 1975 The interrelationships of phalacanthous selachians. Neues Jahrbuch für

Geologie und Paläontologie, Monatshefte, 1975:553–567.

Mustafa, H.A. 2000. Fish teeth from the Upper Umm Ghudran Formation (Late Santonian) of

NW-Jordan. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte, 2000:595–

612.

Mustafa, H.A., G.R. Case, and I.S. Zalmout. 2002. A new selachian fauna from the Wadi Umm

Ghudran Formation (Late Cretaceous) - Central Jordan. Neues Jahrbuch für Geologie und

Paläontologie, Abhandlungen, 226:419–444.

Prowell D.C., R.A. Christopher R.A., K.E. Waters, and S.K. Nix, 2003. The chrono- and

lithostratigraphic significance of the type section of the Middendorf Formation, Chesterfield

County, South Carolina. Southeast Geology, 42:47–66.

Roemer F. 1849. Texas. Mit besonderer Ru¨cksicht auf deutsche Auswanderung und die physischen

Verha¨ltnisse des Landes nach eigener Beobachtung geschildert. Adoph Marcus, Bonn. p. 1– 464

Romer, A.S. 1942. Notes on certain American Paleozoic fishes. American Journal of Science,

240:216–228.

Shimada, K. and D.J. Cicimurri. 2006. The oldest record of the Late Cretaceous anacoracid

shark, Squalicorax pristodontus (Agassiz), from the Western Interior, with comments on

Squalicorax phylogeny. New Mexico Museum of Natural History and Science, Bulletin

35:177–184.

Shimada, K. and D.J. Martin. 2008. Fossil fishes from the basal Greenhorn Limestone (Upper

Cretaceous: late Cenomanian) in Russell County, Kansas. Fort Hays Studies, 2:89–103.

Siverson, M. 1992. Biology, Dental Morphology and Taxonomy of Lamniform Sharks from the

Campanian of the Kristianstad Basin, Sweden. Palaeontology, 35:519–554. 21 Siverson, M. 1996. Lamniform sharks of the mid Cretaceous Alinga Formation and Beedagong

Claystone, Western Australia. Palaeontology, 39: 813–849.

Siverson, M. 1997. Sharks from the mid-Cretaceous Gearle Siltstone, southern Carnarvon Basin,

Western Australia. Journal of Vertebrate Paleontology, 17: 453–465.

Sohl N.F. and Owens J.P., 1991. Cretaceous stratigraphy of the Carolina coastal plain. Pp. 191–

20 in N.F. Sohl and J.P. Owens (eds). The Geology of the Carolinas: Carolina Geological

Society 50th Anniversary Volume. Knoxville, TN: University of Tennessee Press.

Sørensen, A.M., F. Surlyk, J. Lindgren. 2013. Food resources and habitat selection of a diverse

vertebrate fauna from the upper lower Campanian of the Kristianstad Basin, southern

Sweden. Cretaceous Research, 42:85–92.

Underwood, C.J. and S.L. Cumbaa. 2010. Chondrichthyans from a Cenomanian (Late

Cretaceous) bonebed, Saskatchewan, Canada. Palaeontology, 53:903–944.

Vullo, R. 2005. Selachians from the type Campanian area (Late Cretaceous), Charentes,

western France. Cretaceous Research, 26:609–632.

Vullo, R. 2007. Discovery of the genus Proheterodontus Underwood and Ward. (Elasmobranchii,

Neoselachii) in the Upper Jurassic of southwestern France. Neues Jahrbuch für Geologie

und Paläontologie, Abhandlungen, 243:371–375.

Welton, B.J. and R.F. Farish. 1993. The Collector’s Guide to Fossil Sharks and Rays from the

Cretatceous of Texas. Lewisville, TX: Before Time.

Whitley, G.P. 1939. Taxonomic notes on sharks and rays. Australian Zoologist, 9:227–262.

Woodward, A.S. 1886. On the palaeontology of the selachian genus Notidanus Cuvier.

Geological Magazine, 3:205–217. 22 Woodward, A.S. 1889. Sclerorhynchus atavus. Remarks. Proceedings of the Zoological Society

of London, 1889:449–451

Zalmont, I.S. and H.A. Mustafa. 2001. A selachian fauna from the Late Cretaceous of Jordan.

Abhath Al–Yamouk Basic Sciences and Engineering, 10:374–434. 23

Academic Vita Taylor A. Kightlinger

EDUCATION Penn State Erie, The Behrend College Graduating May 2017 Bachelor of Science in General Biology Behrend Honors College Completed Schreyer Honors College In progress Dean’s List Every semester

WORK EXPERIENCE Veterinary Technician at Animal Kingdom Pet Hospital (2015 - present) . Perform diagnostics through bloodwork, radiology, microscopy, bacterial & fungal cultures . Assist doctors and monitor patients during surgeries, dental procedures, and diagnostics . Administer medications to both out-patients and hospitalized pets . Execute preemptive healthcare measures through vaccinations, insect preventions, and proper grooming . Sanitize work stations, exam rooms, equipment, surgical clothing and tools

Behrend Work Study - Preparatory Lab Behrend Work Study - Chemistry Lab Grader Technician (2013 - 2016) (2015) . Prepared all materials and equipment needed . Hand-chosen by the instructor to grade for biology labs chemistry laboratory reports . Operated autoclaves, fume hoods, and hot Aldi - Associate & Cashier (2014 - 2015) plates . Handled large amounts of money per shift . Assembled various types of Petri dishes and . Operated a pallet jack and a box baler other solutions Arby’s (2012 - 2015) . Utilized top-loader scales, analytical balances, . Trained in cashier, drive-thru, kitchen, food pipets, etc. preparation, maintenance, opening, and closing positions . Excelled in custom service

LEADERSHIP/VOLUNTEER EXPERIENCE Publicity Chair & General Member of Reality Laboratory Teaching Assistant (2014 - 2016) Check (2013-2015) Biology Concepts & Biodiversity . Volunteered around campus and in Erie Biology Function & Development of Organisms . Created awareness of our organization via Biology Populations and Communities posters, flyers, and social media . Guided students during experiments . Attended the Alternative Spring Break trip to . Fulfilled demonstrations New Orleans to help reconstruct houses and . Graded assignments restore community gardens that were damaged by Hurricane Katrina

ACHIEVEMENTS/AWARDS STEM Scholar First-year Writing Award Penn State Anonymous Scholarship Award Merwin Scholarship

OTHER ACADEMIC EXPERIENCE Shark Teeth Morphology & Paleontology Drawing & Painting Theatre Production Animal Dissections Ceramics & Sculpture Cinema