USF Libraries
USF Digital Collections

The biomechanics and evolution of shark teeth

MISSING IMAGE

Material Information

Title:
The biomechanics and evolution of shark teeth
Physical Description:
Book
Language:
English
Creator:
Whitenack, Lisa Beth
Publisher:
University of South Florida
Place of Publication:
Tampa, Fla
Publication Date:

Subjects

Subjects / Keywords:
Elasmobranchii
Functional morphology
Feeding
Performance
Finite element analysis
Dissertations, Academic -- Biology -- Doctoral -- USF   ( lcsh )
Genre:
non-fiction   ( marcgt )

Notes

Abstract:
ABSTRACT: Measuring the effects of morphology on performance, and performance on fitness, is necessary to gain a full picture of selection, adaptation, ecology, and evolution. The performance of an organism's feeding apparatus, of which teeth are an integral part, has obvious implications for its fitness and survival. Extant shark teeth encompass a wide variety of shapes, and are often ascribed qualitative functions without any biomechanical testing, employing terminology such as gripping, piercing, crushing, cutting, or tearing. Additionally, teeth also comprise the vast majority of the fossil record of sharks. Therefore to understand the evolution of the shark feeding mechanism, we must understand the contribution of all parts of the feeding apparatus, including the teeth. Performance testing of extant and extinct shark teeth, nanoindentation of shark teeth, finite element analysis of tooth morphology, and phylogenetically informed analyses of shark tooth morphology and ecology were employed to elucidate the relationship between performance, ecology, and evolution. Performance testing of teeth in puncture and draw revealed few morphological patterns, indicating that most morphologies are functionally equivalent. Finite element modeling of teeth in puncture, draw, and holding showed that shark teeth are structurally strong and unlikely to fail during feeding. Evolutionary analyses of tooth shape and ecology showed no relationship between morphology, habitat, and diet. These results have significant implications for the shark paleontology, where the shapes of shark teeth are used to make assumptions about ecology and evolution.
Thesis:
Dissertation (Ph.D.)--University of South Florida, 2008.
Bibliography:
Includes bibliographical references.
System Details:
Mode of access: World Wide Web.
System Details:
System requirements: World Wide Web browser and PDF reader.
Statement of Responsibility:
by Lisa Beth Whitenack.
General Note:
Title from PDF of title page.
General Note:
Document formatted into pages; contains 354 pages.
General Note:
Includes vita.

Record Information

Source Institution:
University of South Florida Library
Holding Location:
University of South Florida
Rights Management:
All applicable rights reserved by the source institution and holding location.
Resource Identifier:
aleph - 002046385
oclc - 496011736
usfldc doi - E14-SFE0002678
usfldc handle - e14.2678
System ID:
SFS0026995:00001


This item is only available as the following downloads:


Full Text
xml version 1.0 encoding UTF-8 standalone no
record xmlns http:www.loc.govMARC21slim xmlns:xsi http:www.w3.org2001XMLSchema-instance xsi:schemaLocation http:www.loc.govstandardsmarcxmlschemaMARC21slim.xsd
leader nam 2200409Ka 4500
controlfield tag 001 002046385
005 20100106135207.0
007 cr mnu|||uuuuu
008 100106s2008 flu s 000 0 eng d
datafield ind1 8 ind2 024
subfield code a E14-SFE0002678
035
(OCoLC)496011736
040
FHM
c FHM
049
FHMM
090
QH307.2 (Online)
1 100
Whitenack, Lisa Beth.
4 245
The biomechanics and evolution of shark teeth
h [electronic resource] /
by Lisa Beth Whitenack.
260
[Tampa, Fla] :
b University of South Florida,
2008.
500
Title from PDF of title page.
Document formatted into pages; contains 354 pages.
Includes vita.
502
Dissertation (Ph.D.)--University of South Florida, 2008.
504
Includes bibliographical references.
516
Text (Electronic dissertation) in PDF format.
3 520
ABSTRACT: Measuring the effects of morphology on performance, and performance on fitness, is necessary to gain a full picture of selection, adaptation, ecology, and evolution. The performance of an organism's feeding apparatus, of which teeth are an integral part, has obvious implications for its fitness and survival. Extant shark teeth encompass a wide variety of shapes, and are often ascribed qualitative functions without any biomechanical testing, employing terminology such as gripping, piercing, crushing, cutting, or tearing. Additionally, teeth also comprise the vast majority of the fossil record of sharks. Therefore to understand the evolution of the shark feeding mechanism, we must understand the contribution of all parts of the feeding apparatus, including the teeth. Performance testing of extant and extinct shark teeth, nanoindentation of shark teeth, finite element analysis of tooth morphology, and phylogenetically informed analyses of shark tooth morphology and ecology were employed to elucidate the relationship between performance, ecology, and evolution. Performance testing of teeth in puncture and draw revealed few morphological patterns, indicating that most morphologies are functionally equivalent. Finite element modeling of teeth in puncture, draw, and holding showed that shark teeth are structurally strong and unlikely to fail during feeding. Evolutionary analyses of tooth shape and ecology showed no relationship between morphology, habitat, and diet. These results have significant implications for the shark paleontology, where the shapes of shark teeth are used to make assumptions about ecology and evolution.
538
Mode of access: World Wide Web.
System requirements: World Wide Web browser and PDF reader.
590
Co-advisor: Philip J. Motta, Ph.D.
Co-advisor: Daniel C. Simkins, Jr., Ph.D.
653
Elasmobranchii
Functional morphology
Feeding
Performance
Finite element analysis
0 690
Dissertations, Academic
z USF
x Biology
Doctoral.
773
t USF Electronic Theses and Dissertations.
856
u http://digital.lib.usf.edu/?e14.2678



PAGE 1

The Biomechanics and Evolution of Shark Teeth by Lisa Beth Whitenack A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy Department of Biology College of Arts and Sciences University of South Florida Co-Major Professor: Philip J. Motta, Ph.D. Co-Major Professor: Daniel C. Simkins, Jr., Ph.D. Stephen Deban, Ph.D. Thomas Koob, Ph.D. James Garey, Ph.D. Carl Luer, Ph.D. Date of Approval: November 7, 2008 Keywords: Elasmobranchii, functional mor phology, feeding, performance, finite element analysis Copyright 2008 Lisa B. Whitenack

PAGE 2

Dedication I dedicate this work to my grandmother Marie Prokuski. She has been one of my biggest cheerleaders and an inspiration.

PAGE 3

Acknowledgements I am indebted to far more people that I can hope to list here. I first want to thank Phil Motta for being a wonderful mentor, teacher, co ffee enabler, and friend for the last seven years. I am glad that I decided to work on squishy things with you and am grateful for your patience and every opportuni ty that you gave me. Secondly, Scott Kowalke has been my rock through this entire experience. I certainly would not have made it through this journey without his constant support, love, laughter, and encouragement. I owe a huge thank you to my co-advisor Daniel Simkins for his knowledge, guidance, and friendship, and for teaching me correct engineering voca bulary such as extreme fiber distance. Many others have contributed to the work herein, including Drs. Thomas Koob, Robert Hueter, Stephen Deban, James Garey, Carl Lu er, and Florence Thomas. Gordon Hubbell, Jack Morris (Mote Marine Laboratory), Anthony Cornett, Josh Collins (FWRI), Mason Dean, Geremy Cliff (Kwazulu Natal Shar ks Board), James Neurndorf, Hubbards Marina, Kyle Mara and The Living Seas (W alt Disney World) provided specimens. Access to and assistance with collections were provided by Gordon Hubbell, Rob Robins (FLMNH), Andrew Piercy (FLMNH), Irv Quitmeyer (FLMNH), George Burgess (FLMNH), Marianne Rogers (FMNH), Jeff Clayton (NMNH), Jeff Williams (NMNH), Jerry Finan (NMNH), Robert Purdy (NMNH), Scott Schaeffer (AMNH), Barbara Brown (AMNH), Radford Arrindell (AMNH), and John Maisey (AMNH). Thank you to Betsy Dumont, Sean Wearle, Mason Meers, Ray Martinez, Mason Dean, Daniel Huber,

PAGE 4

Theodore Garland, Jr., Mike Melendez, Doug Pr ingle, Dan Hernandez, Adam Summers, Jason Rohr, Earl McCoy, Makoto Harai, As hok Kumar, and Gregory Herbert for their technical support and guidance. An army of volunteers made this research possible, especially Danita Collins, Brianna Fulcher, Sarah Koh, Eric Lambert, Joshua Lipham, Alison Meyers, Janne Pfeiffenberger, Courtney Phillips, Stacy Villanueva, and my mom Karen Whitenack. Alpa Wintzer, Angela Coll ins, Laura Habegger, Samantha Mulvany, Daniel Huber, Mason Dean, and Kyle Mara prov ided valuable discussion and stress relief in the lab. Funding for this project was provi ded by the American Elasmobranch Society, Geological Society of America, Porter Family Foundation, IGERT Program Grant DGE 0021681, Society for Integrative and Comp arative Biology, Tampa Bay Fossil Club, Sarah Spector, University of South Florida Department of Biology, University of South Florida Foundation, and University of South Florida Office of Research.

PAGE 5

i Table of Contents List of Tables iii List of Figures iv Abstract vi Introduction 1 Performance of Shark Teeth During Punctu re and Draw: Implications for the Mechanics of Cutting 12 Abstract 12 Introduction 13 Materials and Methods 17 Specimens for Performance Testing 17 Performance Testing 19 Tooth Morphometrics 21 Statistical Analyses 22 Results 24 Puncture 24 Unidirectional Draw 26 Phylogenetic & Morphometric Relationships 27 Discussion 27 Biomechanics of Prey Items 28 Biomechanics of Extant Teeth 30 Fossil Tooth Mechanics 36 Conclusions 39 Biology Meets Engineering: The Structural Me chanics of Shark Teeth 52 Abstract 52 Introduction 53 Shark Tooth Materials 56 Materials and Methods 57 Nanoindentation 57 Finite Element Analysis 60 Specimens 60 Modeling 60 Results 62 Nanoindentation 62

PAGE 6

ii Finite Element Analysis 63 Discussion 64 Nanoindentation 65 Finite Element Analysis 68 Conclusions 72 Evolutionary Relationships Between Shark Tooth Morphology and Ecology 82 Abstract 82 Introduction 83 Materials and Methods 86 Specimens 86 Ecological Information 86 Tooth Morphometrics 87 Statistical Analysis 89 Results 90 Habit and Depth 90 Diet 91 Discussion 93 Conclusions 100 Conclusions 108 References 113 Appendices 143 Appendix A 143 Appendix B 146 Appendix C 220 Appendix D 228 Appendix E 241 Appendix F 246 Appendix G 335 Appendix H 342 About the Author End Page

PAGE 7

iii List of Tables Table 1.1 Tooth and jaw position informa tion for each extant species in this study. 45 Table 1.2 Means for force to puncture (F p ) (N = Newtons) + standard error. 46 Table 1.3 Means for maximum force during puncture (F max ) (N = Newtons) + standard error. 47 Table 1.4 Means for energy to puncture (E p ) (N/mm) + standard error. 48 Table 1.5 Two-way ANOVA results for pooled species and pooled prey. 49 Table 1.6 One-way ANOVA results for species within prey items. 50 Table 1.7 Means for maximum force during draw (F draw ) (N = Newtons) + standard error. 51 Table 2.1 Tooth and jaw position information for each extant species in this study. 80 Table 2.2 Material properties of tooth materials for vertebrates. 81

PAGE 8

iv List of Figures Figure 1.1 Phylogenetic tree of species included in this study and tooth morphology, based on Compag no, 1988; Martin et al., 1992; Naylor, 1992; Shirai, 1996; Gr ogan and Lund, 2004. 41 Figure 1.2 Tooth measurements taken for canon ical correlation analysis. 42 Figure 1.3 Force-displacement trace for puncture of Haemulon plumieri by an anterior Isurus oxyrinchus tooth. 43 Figure 1.4 Force-displacement trace for unidirectional draw through Haemulon plumieri by an anterior Isurus oxyrinchus tooth. 44 Figure 2.1 Testing sites for nanoindentati on on a cross-section of a generalized tooth cusp. 74 Figure 2.2 Phylogenetic tree of species included in this study, based on Compagno, 1988; Martin et al., 1992; Naylor, 1992; Shirai, 1996; Grogan and Lund, 2004. 75 Figure 2.3 Loading regimes illustrated on bull shark tooth ( Carcharhinus lecuas, left) and lateral view of shortfin mako shark tooth ( Isurus oxyrinchus right). 76 Figure 2.4 Representative finite element models (FEMs) loaded in puncture. 77 Figure 2.5 Representative FEMs loaded in draw on the distal cutting edge. 78 Figure 2.6 Representative FEMs loaded in holding on the lingual face of the tooth. 79 Figure 3.1 Phylogenetic tree of species included in this study, based on Compagno, 1988; Martin et al ., 1992; Naylor, 1992; Shirai, 1996; Goto, 2001. 101 Figure 3.2 Phylogenetic tree of species included in this study, with tooth morphology and ecology. 102

PAGE 9

v Figure 3.3 Upper jaw of Carcharhinus leucas illustrating tooth positions sampled and base overlap measurement (BO). 106 Figure 3.4 Tooth measurements taken for e volutionary analysis 107

PAGE 10

vi The Biomechanics and E volution of Shark Teeth Lisa Beth Whitenack ABSTRACT Measuring the effects of morphology on performance, and performance on fitness, is necessary to gain a full picture of selection, adaptation, ecology, and evolution. The performance of an organisms feeding appa ratus, of which teeth are an integral part, has obvious implications for its fitness and survival. Extant shark teeth encompass a wide variety of shapes, and are often as cribed qualitative f unctions without any biomechanical testing, employing terminol ogy such as gripping, piercing, crushing, cutting, or tearing. Additionally, teeth also comp rise the vast majority of the fossil record of sharks. Therefore to understand the evolution of the sh ark feeding mechanism, we must understand the contribution of all parts of the feeding a pparatus, including the teeth. Performance testing of extant and extinct shark teeth, nanoindent ation of shark teeth, finite element analysis of tooth morphology, and phylogenetically informed analyses of shark tooth morphology and ecology were employed to elucidate the relationship between performance, ecology, and evolution. Performance testing of teeth in puncture and draw revealed few morphological patterns, indicating that most morphologies are functionally equivalent. Finite element modeling of teeth in puncture, draw, and holding showed that shark teeth are structurally strong and unlikely to fail during feeding.

PAGE 11

vii Evolutionary analyses of tooth shape a nd ecology showed no relationship between morphology, habitat, and diet. These results ha ve significant implications for the shark paleontology, where the shapes of shark te eth are used to make assumptions about ecology and evolution.

PAGE 12

Introduction Functional morphology seeks to explain the relationship between form, that is the shape of the feature in question, and function, how a feature works or what it does (Bock and von Walhert, 1965; Dullemeijer and Barel, 1977). The exploration of this relationship can be used to infer biological role (how the feature is used in the context of its environment) and adaptive scenarios, which in turn become starting points for other evolutionary questions, such as the origin or evolutionary consequences of the features in question (Plotnick and Baumiller, 2000). Biomechanics, the application of quantitative engineering techniques to determine how organisms perform mechanical functions, the design of morphological systems, and the relationship of these to the organisms environment, is one way to explore functional morphology (Lauder, 1991; Biewiner, 1992; Koehl, 1996). Biomechanics allows functional properties of biological structures and the ability of an organism to carry out these functions (performance) to be quantified and their effects on biological role to be tested (Koehl, 1996; Plotnick and Baumiller, 2000). Measuring the links between morphology, performance, and fitness is necessary to gain a full picture of selection and adaptation (Bock, 1980; Arnold, 1983; Wainwright, 1996). A number of recent studies of vertebrate functional morphology have specifically addressed feeding, as the performance of an organisms feeding apparatus has obvious implications for its fitness and survival (Biknevicius et al., 1996; Evans and Sanson, 1

PAGE 13

1998; Deufel and Cundall, 1999; Schwenk, 2000; Dumont et al., 2005; Jones, 2008). Though many of functional morphological studies on feeding focus on osteichthyans (Lauder, 1980; Bemis and Lauder, 1986; Wainwright, 1988; Dutta, 1992; Sanford, 2001; Konow and Sanford, 2008), there are a growing number that incorporate chondrichthyans (Springer, 1961; Moss, 1977; Frazzetta, 1994; Motta, 2004; Dean et al., 2008; Motta et al., 2008). The majority of studies of elasmobranchs concern muscle function and the movement of cranial components, while the role of teeth has largely been ignored despite the fact that they are an important part of the feeding apparatus (but see Ramsay, 2007; Dean, 2008). The cartilaginous nature of the skeleton and continuous tooth replacement have lead to a chondrichthyan fossil record composed primarily of teeth. This makes fossilzed teeth our primary tool for studying shark evolution through the over 400 million years of their existence. When exploring functional morphology in extant animals, not only can the structure of the organ in question be analyzed, but also behavior and physiology can be observed to support hypotheses made based on structure. However, behavior cannot be directly fossilized and is lost when studying the functional morphology of extinct animals. This limits the investigator, who may only able to use structure and paleoenvironmental data to reconstruct function (Reif, 1983). Even so, the precision with which function can be inferred is limited by the degree of structural information available (Plotnick and Baumiller, 2000). Therefore, studying the functional morphology of shark teeth not only elucidates the biological role that teeth play in feeding, but also provides insight specifically into the evolution of shark feeding as teeth are often the only structures available in the fossil record (Carroll, 1988). 2

PAGE 14

Ecomorphological studies investigate the interactions of morphology, ecology, behavior, and performance (Williams, 1972; Karr and James, 1975; Bock, 1994). Due to the close relationship between these parameters, it is assumed that evolution of morphology and ecology are tightly correlated (Losos, 1990), resulting in the predictive power of one to the other (Weins and Rotenberry, 1980; Motta and Kotrschal, 1992). Shark tooth morphotypes and their presumed function present an interesting test of this ecomorphological paradigm. Extant shark teeth encompass a wide variety of shapes, including teeth with triangular serrated cusps, oblique serrated and non-serrated cusps, notched serrated cusps, non-serrated recurved cusps, multicusped teeth, and flattened tooth pavements. These teeth perform a variety of biological functions including grasping, cutting and crushing, though performance testing has not been applied to shark teeth. Consequently, tooth forms are often ascribed qualitative functions without any biomechanical testing (Cappetta, 1986, 1987; Motta, 2004). Small teeth with lateral cusplets are characterized as clutching-type. These are found within the Orectolobidae, Ginglymostomidae, Squatinidae, and Scyliorhinidae. Tearing-type teeth have narrow, tall cusps and are usually not serrated; examples include teeth of the lamniform Isurus and Mitsukurina species. Teeth whose crowns are lingo-labially flattened and widen towards the base are cutting-type; many of the Carcharhinidae fall into this category. Molariform teeth, such as those found in the heterodontids and many batoids, fall into crushing-type or grinding-type (Moss, 1977; Cappetta, 1987; Frazzetta, 1988). These categories are by no means exclusive. Heterodontus have clutching teeth anteriorly and crushing teeth posteriorly, while Isurus has tearing teeth anteriorly and cutting teeth laterally (Cappetta, 1987). 3

PAGE 15

Inevitably, links between these functional morphotypes and ecology are made. For example, sharks with clutching teeth are often benthic or benthopelagic and presumably seize elusive midwater prey. Elasmobranchs with crushing dentition tend to prey upon hard prey such as bivalves and mollusks, as well as small fishes and cephalopods; they tend to be benthic or benthopelagic (Cappetta, 1987). Those with grinding dentitions are usually benthic and also feed upon hard prey (Reif, 1976; Nobiling, 1977; Cappetta, 1987; Summers et al., 2004). These relationships lead to predictions about ecology for taxa where information about diet and habitat are lacking. For example, Chlamydoselachus anguineus, the frilled shark, is a poorly known deepwater shark. It has been predicted that this shark feeds on cephalopods and bottom associated fishes based on its tooth morphology (Compagno, 2001). Fossil sharks have exhibited a wide diversity of tooth morphologies, many of which are not found in extant forms. Given the abundance of teeth in the fossil record, compared to other parts of the body, tooth morphology has been used to divide sharks into different evolutionary levels: cladodont, hybodont, and modern. These levels are also supported by other characteristics, such as jaw and fin structure (Schaeffer, 1967). These levels reflect general trends and do not imply specific evolutionary relationships. Zangerl (1981) lists a number of species with cladodont-level tooth morphologies that are not closely related to each other, based on non-dental characters. The cladodont level represents the earliest sharks, found mainly from the Middle Devonian (391 mya) to the Pennsylvanian (323 mya). Most of the genera included in this level possess cladodont teeth. This morphology has a large medial cusp with numerous lateral cusplets on each side, with a flattened disk-like base (Schaeffer, 1967). The 4

PAGE 16

xenacanth tooth morphology is also included in the cladodont level, though it does not appear until the Early Carboniferous (Schaeffer, 1967; Benton, 1997). This morphology is best illustrated by Xenacanthus. The medial cusp is the smallest, with larger lateral cusps on each side (Schaefer, 1967). The hybodont level sharks may have existed as early as the Devonian, but were the dominant group from the Triassic (248 mya) to the Jurassic (206 mya) (Schaeffer, 1967; Maisey, 1982). The morphology of the teeth of hybodont level sharks was much less generalized than those of the cladodont level. Tooth morphology is either multicusped, though in a less dramatic fashion than the cladodonts, or molar-like. The medial cusp in the multicusped teeth varies in height from much taller to almost the same height as the lateral cusplets (Cappetta, 1987). Modern elasmobranchs first appear in the late Paleozoic and became the dominant group during the Cretaceous (144 to 65 mya) (Benton, 1997). Due to the cartilaginous nature of the skeleton, the same tendency to predict ecology and function from tooth morphology exists for fossil chondrichthyan teeth (Peyer, 1968; Cappetta, 1987; Williamson et al., 1993; Cicimurri, 2000, 2004; Stahl and Parris, 2004). For example, the Jurrasic shark Sphendous has high crowned teeth (tearing type) and has been hypothesized to eat soft-bodied invertebrates, while cochliodont holocephalans are hypothesized to be durophagous based on their molariform dentition (Peyer, 1968; Cappetta, 1987). Inferences about shark evolution are also often made based on tooth morphology (Schaeffer, 1967; Maisey, 1982). For example, it has been hypothesized that selection for larger bladed teeth occurred with a change to a macrophagous diet (Williams, 2001). 5

PAGE 17

Tooth function The vast majority of tooth function studies have been performed on mammals (e.g. Churcher, 1985; Van Valkenburgh and Ruff, 1987; Van Valkenburgh, 1988; Freeman, 1992; Lucas et al., 1994; Biknevicius et al., 1996; Evans and Sanson, 1998; Fenton et al., 1998; Evans and Sanson, 2005; Freeman and Lemen, 2007). While a large number of functional morphological studies of teeth address osteichthyans (e.g. Wainwright, 1987; Kotrshal and Goldschmidt, 1992; Hernandez and Motta, 1997; Wautier et al., 2001), comparatively few studies exist for shark teeth (Nobiling, 1977; Frazzetta, 1988; Powlik, 1995; Ramsay and Wilga, 2007; Dean et al., 2008). Frazzetta (1988) was the first to specifically address the biomechanics of cutting in shark teeth and its relation to form. Applying blades and teeth from various shark species to compliant materials, he observed that smooth, slender teeth, such as those of the mako shark Isurus oxyrinchus, appeared better suited for puncturing and piercing. The recurved tips of these teeth could possibly enhance the probability of initial prey penetration, as hypothesized for snakes with similar teeth (Frazzetta, 1966; Cundall and Deufel, 1999; Deufel and Cundall, 1999). Serrated teeth, such as those of the white shark Carcharodon carcharias and tiger shark Galeocerdo cuvier, appeared to be more useful for slicing and cutting, though are more susceptible to binding of the teeth in prey than the smooth, non-serrated teeth. Though the majority of his study was on teeth of carnivorous dinosaurs, Abler (1992) included fossil teeth from Carcharodon sp. He qualitatively tested three hypotheses about the mechanics of cutting for smooth and serrated teeth and found that 6

PAGE 18

serrated blades trap material between serrations and thus rip tissues as they are carried by the tooth. Smooth blades concentrate a large force on the cutting edge, creating high pressure that crushes the material beneath it, producing the cut. Results also indicated that the act of drawing the tooth across the substrate has different functions with respect to cut propagation, depending on whether the tooth is smooth or serrated. In serrated teeth, the draw provides the motion required by the grip and rip hypothesis. In smooth teeth, the draw breaks the frictional grip between the blade and substrate, allowing all of the applied downward force to rest on the cutting edge. Several studies have addressed the orientation of teeth with respect to the jaw and its role during feeding. One study suggests that the inward turned tips of the anterior teeth of the white shark Carcharodon carcharias may make gouging and grasping of prey more efficient (Powlik, 1995). It is also hypothesized that the more labially inclined upper anterior teeth of the sand tiger shark Carcharias taurus puncture prey, while the more lingually inclined lower anterior teeth may be used for initial prey grasping (Lucifora et al., 2001). More recent studies focus on reorientation of teeth during feeding and their dual roles in gripping and either crushing or protecting the jaw as it hits the seafloor (battering) (Ramsay and Wilga, 2007; Dean et al., 2008). In general, the biomechanics of the tooth itself have been ignored, including the structural mechanics of the tooth. Chondrichthyans are capable of producing a range of bite forces. Static equilibrium models calculate the average anterior bite force for Etmopterus spinax to be 1.01 N (mean SL = 32.5 cm) (Claes & Malefet, unpublished data), while a bite force of 19.6 N was calculated for a 45.1 cm SL Squalus acanthias (Huber and Motta, 2004). The 7

PAGE 19

highest calculated bite forces are well over 1000 N. Static equilibrium models estimate the posterior theoretical bite force of the great hammerhead Sphyrna mokarran (4.3 m TL) to be 6080 N (Mara et al., unpublished data); finite element models of a 6.4 m TL white shark, Carcharodon carcharias, estimate bite forces of 9320 N anteriorly and 18216 N posteriorly (Wroe et al., 2008). As teeth are subjected to these sometimes extreme loads during feeding, they may undergo stress, strain, and potentially failure. Teeth do not heal; to continue performing their biological role, teeth must resist breakage until they are shed. While performance is certainly subject to natural selection (Arnold, 1983; Bennet, 1991), attributes related to structural strength such as material properties and overall shape may also be subjected to natural selection (Erickson et al., 2002; Lucas et al., 2008). Therefore, both prey processing ability and structural parameters must be considered to understand the evolution of shark teeth. Shark tooth materials Shark teeth can be divided into two zones: the crown and the root or base. Shark teeth are also composite material structures, with two distinct structural components: a central core of dentine covered by enameloid, an enamel-like substance formed from both odontoblasts and ameloblasts (Poole, 1967a). Some elasmobranchs also have a central pulp cavity (orthodont teeth), while in others the root extends into the central core of the crown (osteodont teeth) (Cappetta, 1987; Compagno, 1988). Enameloid is a highly mineralized tissue, composed primarily of hydroxyapatite crystallites. These crystallites are arranged in bundles that vary in orientation depending on the location within the tooth (Poole, 1967a; Gillis and Donoghue, 2007). In modern 8

PAGE 20

sharks, these bundles tend to align into three layers: a thin superficial layer with randomly arranged single cystallites (shiny layer enameloid, SLE), followed by a parallel-fibered layer (PFE), and an innermost tangled-fiber layer (TFE) (Gillis and Donoghue, 2007). Previous work has shown that the PFE imparts tensile strength, while the TFE provides resistance to compressive forces (Preuschoft et al., 1974). Together, shark enameloid forms a layer that is 0.2-0.9 mm thick (Preuschoft et al., 1974). While Devonian cladodont sharks lacked enameloid on their teeth, Mississippian cladodont teeth, as well as xenacanthids, have a thin enameloid layer (Dean, 1909; Goto, 1991). Hybodont sharks with multicuspid teeth also have a thin layer of enameloid, but those with molariform teeth do not (Goto, 1991). Elasmobranch dentine is composed of mineralized collagenous tissue (Bradford, 1967; Johansen, 1967). Two types of dentine occur in shark teeth. Osteodentine superficially resembles spongy bone; the dentine surrounds vascular canals, similar to osteons. Orthodentine does not contain dental osteons. Instead it contains smaller parallel branching tubules that provide a banded appearance (Compagno, 1988). Osteodentine forms the base in all shark teeth. The crown can be composed primarily of either orthodentine (orthodont), such as the teeth of carcharhiniform sharks, or osteodentine (osteodont), as in lamniform sharks (Mertinene, 1982; Compagno, 1988). Cladodont and hybodont teeth tend to be composed of both orthodentine and osteodentine, while xenacanthid teeth are composed only of orthodentine (Goto, 1991). Despite a prolific data base on mammalian tooth properties, especially humans (e.g. Waters, 1980; Brear et al., 1990), the material properties of shark teeth and their components have not been studied to date. 9

PAGE 21

Goals and hypotheses The overall goal of this study was to investigate the biomechanics and evolution of extant and extinct shark teeth. All parts of this study revolve around tooth morphology, as tooth forms are often ascribed qualitative functions without any biomechanical testing, and morphology is used to infer function and ecology in both extant and extinct sharks. For the first part of this study, I measured the performance of teeth in puncture and unidirectional draw for ten species of extant shark and three extinct shark clades during puncture and unidirectional draw (cutting). The hypothesis for this portion of the study is that performance is determined by tooth morphology. The specific goals of this portion of the study were to: (1) Determine the forces necessary for individual teeth to penetrate a variety of fish and crustacean prey representative of shark diets; (2) Determine what differences in penetration force and efficiency occur among tooth types; (3) Compare performance between different cutting regimes for a given tooth morphology and (4) Determine which morphological aspects, if any, of tooth shape are predictive of tooth performance. These results have implications for studies of fossil chondrichthyans, where typically tooth morphology is used to predict feeding ecology in the absence of behavioral data (Peyer, 1968; Zangerl, 1981; Lund, 1985, 1990; Whitenack et al., 2002; Elliott et al., 2004). In the second part of the study, the structural mechanics of fossil and extant shark teeth were investigated. The first goal was to determine the material properties for enameloid, osteodentine, and orthodentine via nanoindentation. The hardness and elastic modulus of these tissues is not known, and is necessary for the second portion of this 10

PAGE 22

study. I then used finite element analysis to visualize stress distributions of fossil and extant shark teeth during puncture, unidirectional draw (cutting), and holding. The specific goals for this portion of the study were to determine if tooth morphologies are more structurally strong during one loading regime versus another and to examine the role of morphological features, such as notches or cusp shape, on stress distribution. For the last part of the study, I employed phylogenetic comparative methods to test, within an evolutionary context, whether a relationship exists between shark tooth morphology and ecology. While connections between shark tooth morphology, ecology, and evolution are often cited, the existence of a firm relationship between tooth morphology and ecology has not been rigorously tested. Based on ecomorphological principles, I hypothesize that diet and habitat are predictive of shark tooth shape. 11

PAGE 23

Chapter 1: Performance of Shark Teeth during Puncture and Draw: Implications for the Mechanics of Cutting Abstract The performance of an organisms feeding apparatus has obvious implications for its fitness and survival. However, the majority of studies that focus on chondricthyan feeding tend to address muscle function and the movement of cranial components, while the role of teeth has largely been ignored, despite the fact that they shark teeth vary greatly in morphology and are instrumental in prey procurement and processing. Studying the functional morphology of shark teeth not only elucidates the biological role that teeth play in feeding, but also provides insight specifically into the evolution of shark and vertebrate feeding. In this study, I investigate the performance of two general categories of extant teeth, tearing-type and cutting-type, as well as three fossil morphologies. The goals of this study are to: (1) Determine the forces necessary for individual teeth to penetrate a variety of fish and crustacean prey representative of shark diets; (2) Determine what differences in penetration force and efficiency occur among tooth types; (3) Compare performance between different cutting regimes for a given tooth morphology and (4) Determine which morphological aspects, if any, of tooth shape are predictive of tooth performance. To determine the loads experienced by shark teeth during puncture and unidirectional draw, teeth from ten extant shark species and three 12

PAGE 24

fossil species were tested with a MTS MiniBionix II universal testing system. Puncture forces were determined by driving teeth into a variety of prey items (teleost, elasmobranch, and crab) at 400 mm/s. For unidirectional draw, teeth were embedded in a teleost prey item and drawn in parallel at 400 mm/s. Differences in puncturing performance occurred among different prey items, indicating that not all soft prey items are alike. The majority of teeth were able to puncture different prey items, and differences in puncture performance also occurred among tooth types; however, few patterns emerged. Force to puncture was less than the maximum force that occurred during draw tests, however there were no differences between the maximum draw forces and maximum puncture forces. Few morphological patterns were identified. In some cases, broader triangular teeth were less effective at puncturing than narrow-cusped teeth. Teeth from Galeocerdo cuvier, Prionace glauca,Hexanchus griseus, and Sphyrna mokarran were unable to puncture many of soft prey items. The flat surface of the tooth-prey contact may decrease stress on the prey item to the extent that puncture (failure) is not possible. No morphological characteristics were correlated with maximum draw force. Many of the shark teeth in this study were not only able to perform draw and puncture equally well, but tooth morphologies were functionally equivalent to each other. This does not support the use of tooth morphology to predict biological role. Introduction Measuring the effects, or lack thereof, of morphology on performance, and performance on fitness, is necessary to gain a full picture of selection, adaptation, ecology, and evolution (Bock, 1980; Arnold, 1983; Wainwright, 1996). The performance of an 13

PAGE 25

organisms feeding apparatus has obvious implications for its fitness and survival. In this vein, a number of recent studies of vertebrate functional morphology have specifically addressed feeding (Biknevicius et al., 1996; Evans and Sanson, 1998; Deufel and Cundall, 1999; Schwenk, 2000; Dumont et al., 2005; Jones, 2008). The functional morphology of feeding in extant fishes has been studied in detail. Though many of these studies focus on osteichthyans (Lauder, 1980; Bemis and Lauder, 1986; Wainwright, 1988; Dutta, 1992; Sanford, 2001; Konow and Sanford, 2008), there are a growing number that focus on chondrichthyans (Springer, 1961; Moss, 1977; Frazzetta, 1994; Motta, 2004; Dean et al., 2008; Motta et al., 2008). However, the majority of studies on elasmobranchs concern muscle function and the movement of cranial components, while the role of teeth has largely been ignored, despite the fact that they are instrumental in prey procurement and processing (but see Ramsay, 2007; Dean, 2008). Studying the functional morphology of shark teeth not only elucidates the biological role that teeth play in feeding, but also provides insight specifically into the evolution of shark and vertebrate feeding, as sharks represent an early offshoot of vertebrate life (Carroll, 1988). The teeth of elasmobranchs (sharks, skates and rays) are characterized by ligamentous attachment to the cartilaginous jaws (Reif, 1982). The teeth are attached to the margins of the mouth by Sharpeys fibers, which run from the dentine to the dermis of the dental lamina, not to the jaw cartilage itself (Moss, 1970). The nature of this attachment gives teeth some degree of lingo-labial flexibility, which may facilitate tooth penetration, allow teeth to be deflected when hard materials are encountered during feeding, or to improve cutting during head shaking (Frazzetta and Prange, 1987; Frazzetta, 1994; Powlik, 1995; Ramsay and Wilga, 2007; Dean et al., 2008). 14

PAGE 26

Extant shark teeth encompass a wide variety of shapes, including teeth with triangular serrated cusps, oblique serrated and non-serrated cusps, notched serrated cusps, non-serrated recurved cusps, multicusped teeth, and flattened tooth pavements. Shark teeth perform a variety of biological functions including grasping, cutting and crushing, however performance testing has never been applied to shark teeth. Consequently, tooth forms are often ascribed qualitative functions without any biomechanical testing (Cappetta, 1986, 1987; Motta, 2004). Small teeth with lateral cusplets are characterized as clutching-type. These are found within the Orectolobidae, Ginglymostomidae, Squatinidae, and Scyliorhinidae. Tearing-type teeth have narrow, tall cusps and are usually not serrated; examples include teeth of the lamniform Isurus and Mitsukurina species. Teeth whose crowns are lingo-labially flattened and widen towards the base are cutting-type; many of the Carcharhinidae fall into this category. Molariform teeth, such as those found in the heterodontids and many batoids, fall into crushing-type or grinding-type (Moss, 1977; Cappetta, 1987; Frazzetta, 1988). These categories are by no means exclusive. Heterodontus have clutching teeth anteriorly and crushing teeth posteriorly, while Isurus has tearing teeth anteriorly and cutting teeth laterally (Cappetta, 1987). Tooth function has been explored in a number of vertebrates; however, the majority of studies have been performed on mammals (e.g. Lucas, 1982; Churcher, 1985; Van Valkenburgh and Ruff, 1987; Van Valkenburgh, 1989; Freeman, 1992; Lucas et al., 1994; Biknevicius et al., 1996; Freeman and Weins, 1997; Evans and Sanson, 1998; Evans and Sanson, 2005; Evans et al., 2007; Freeman and Lemen, 2007). While a large number of functional morphological studies of teeth address osteichthians (e.g. 15

PAGE 27

Wainwright, 1987; Kotrshal and Goldschmidt, 1992; Hernandez and Motta, 1997; Wautier et al., 2001), comparatively few studies exist for shark teeth (Nobiling, 1977; Frazzetta, 1988; Powlik, 1995; Ramsay and Wilga, 2007; Dean et al., 2008). Frazzetta (1988) was the first to specifically address the biomechanics of cutting in shark teeth and its relation to form. Applying metal blades, rubber blades, and teeth from various shark species to compliant materials, he observed that smooth, slender teeth, such as those of the mako shark Isurus oxyrinchus, appeared better suited for puncturing and piercing. The recurved tips of these teeth could possibly enhance the probability of initial prey penetration. Serrated teeth, such as those of the white shark Carcharodon carcharias, appeared to be more useful for slicing and cutting, though are more susceptible to binding of the teeth in prey than the smooth, non-serrated teeth. Though most of his study was on the dinosaur Tyrannosaurus rex, Abler (1992) included fossil teeth from Carcharodon sp. He qualitatively tested three hypotheses about the mechanics of cutting for smooth and serrated teeth and found that serrated blades trap material between serrations and thus rip tissues as they as carried by the tooth. Smooth blades concentrate a large force on the cutting edge, creating high pressure that crushes the material beneath it, producing the cut. In this study, I investigated the performance of two general categories of extant teeth, tearing-type and cutting-type, as well as three fossil morphologies. The goals of this study are to: (1) Determine the forces necessary for individual teeth to penetrate a variety of fish and crustacean prey representative of shark diets; (2) Determine what differences in penetration force and efficiency occur among tooth types; (3) Compare performance between different cutting regimes for a given tooth morphology and (4) 16

PAGE 28

Determine which morphological aspects, if any, of tooth shape are predictive of tooth performance. Here we report the results of performance testing of teeth in puncture and unidirectional draw for ten species of extant shark and three extinct shark clades. These results have implications for studies of fossil chondrichthyans, where typically tooth morphology is used to predict feeding ecology in the absence of behavioral data (Peyer, 1968; Zangerl, 1981; Lund, 1985, 1990; Whitenack et al., 2002; Elliott et al., 2004). Materials and Methods Specimens for performance testing Teeth from ten shark species were chosen to cover a wide range of extant tooth forms, as opposed to taxonomy: Isurus oxyrinchus, Carcharodon carcharias, Sphyrna mokarran, Galeocerdo cuvier, Carcharhinus leucas, C. limbatus, Negaprion brevirostris, Prionace glauca, Scymnodon ringens, and Hexanchus griseus (Table 1.1, Figure 1.1). Teeth of these species can be grouped in two general categories: tearing and cutting morphs (Cappetta 1987). Tearing teeth are described as has having narrow cusps with distinct cutting edges, e.g. the anterior teeth of Isurus oxyrinchus. Cutting teeth can be further divided into two subgroups: cutting subtype, where teeth are lingo-labially flattened, with broad cusps distally inclined toward the back of the jaw. Serrations are thought to further improve cutting performance. Teeth of Carcharodon carcharias, Galeocerdo cuvier, Hexanchus griseus, Carcharhinus leucas,Prionace glauca, Sphyrna mokarran and the lateral teeth of Isurus oxyrinchus are members of this subgroup. The other subgroup is the cutting-clutching subtype. It should be noted that the original classification of Cappetta (1987) for this subtype confounds dignathic heterodonty and 17

PAGE 29

tooth morphology; in the cutting-clutching subtype, teeth of the one jaw are wide and linguo-labially flattened and those of the other jaw have high and narrow cusps. For this study, the C. limbatus and S. ringens teeth are part of the cutting jaw, while teeth of N. brevirostris are part of the clutching jaw. It should be noted that the clutching teeth of N. brevirostris are not of the same morphology as those of the small, multicusped clutching-type exemplified by the Orectolobidae (Cappetta, 1987). From each species, except as noted on Table 1.1, single teeth from 3 sexually mature individuals were removed from the jaw and air dried. Teeth were removed from the non-functional row to ensure that teeth were not worn and therefore optimally sharp prior to performance testing. Single teeth from three fossil clades, representing basic fossil morphologies that are not seen in the modern sharks, were used: cladodont (Cladodus sp., Stephens Museum 1998-1a), xenacanth (Xenacanthus compressus, USNM 182325), and hybodont (Hybodus sp. USNM 14197). To prevent breakage of fossil specimens, aluminum casts were made for testing. The geometry of each tooth was acquired via Phillips Mx8000 high-resolution x-ray computed tomography scanner (slice thickness = 11.7 600 microns). The scans were segmented using VGStudioMax (Volume Graphics GmbH, Germany). This process generates a stereolithography (STL) surface mesh of the three-dimensional geometry from stacked DICOM images acquired by the CT scanner. Each STL was then imported into Geomagic Studio 6 (Geomagic Inc., USA), which was used to rebuild parts of the fossil teeth, as all three specimens were partially damaged by taphonomic processes. The modified tooth models were then created from a plaster-based powder using a three-dimensional Zprinter 310 for rapid prototyping (Z Corporation, Burlington, MA, USA) and used to make aluminum casts. 18

PAGE 30

Sharp cusp apices were added back to Xenacanthus compressus and Cladodus sp. and cutting edges to X. compressus and Hybodus sp. with a rotary tool and fine sandpaper, based on Cappetta (1987) and Zangerl (1981). The fossil casts and extant teeth were potted in PVC couplers with Labstone dental cement (Heraeus Kulzer Inc., NY, USA), then mounted to a MTS 858 MiniBionix II universal testing system (Eden Prairie, MN, USA). Performance Testing Tooth performance was tested by puncture and unidirectional draw. Puncture, which does not involve lateral movement of the tooth through tissue, occurs when a tooth enters a prey item such as during biting. Unidirectional draw involves lateral movement through tissue and may occur, for instance, during head shaking behavior. For puncture, teeth were tested on five prey items representing a variety of scale thicknesses and toughness of prey: a teleost with thick scales (sheepshead, Archosargus probatocephalus, TL = 27.9-39.4 cm, mean scale thickness + SE = 0.185 + 0.007 mm), a teleost with scales of intermediate thickness (white grunt, Haemulon plumieri, TL = 20.3-34.9 cm, mean scale thickness = 0.121 + 0.009 mm), a teleost with thin scales (ladyfish, Elops saurus, TL = 29.8-48.3 cm, mean scale thickness = 0.058 + 0.002 mm), an elasmobranch (bonnethead, Sphyrna tiburo, TL = 58.8-74.8 cm), and a crustacean (blue crab, Callinectes sapidus, inter-molt crabs only, carapace spine-to-spine width = 13.7-19.1 cm). These prey items were chosen as representative prey items with a range of biomechanical properties, rather than inclusion in all shark diets, though most of the sharks included in this study eat a variety of teleost prey items (Compagno, 1984a, b; 19

PAGE 31

Cortes, 1999). Prey items were placed on a wooden stage lined with sandpaper to prevent the prey item from slipping during tests. The stage rested upon a 5 kN load cell to obtain force magnitudes that occur during puncture. Teeth were positioned with the cusp apex contacting the prey item. For the teleost and elasmobranch prey, teeth were tested on the epaxialis musculature to avoid the vertebral column and the proximal pterygiophores. Individual prey items were used for multiple punctures at different locations. For crustacean prey, teeth were positioned at the center of the dorsal carapace and individual crabs were only used once. For each puncture trial, teeth were driven into the prey item at a rate of 400 mm/s. This upper limit of the MTS approximates the rate of movement of the upper jaw of carcharhinid sharks during prey capture (460 mm/s) (Motta and Lowry, unpublished data). Tooth displacements were set minimally to the length of the cusp. Two prey items, E. saurus and S. tiburo, deformed greatly before tooth penetration, necessitating displacements to be set to values greater than cusp length. A high speed Redlake PCI 500 or 1000 Motionscope digital video camera (Tallahassee, FL, USA) at 500 fields per second allowed synchronization of initial penetration of the tooth and the force as recorded by the MTS. Three trials were recorded for each tooth. From each puncture trial, force at initial puncture (F p ) and maximum puncture force (F max ) were measured. Energy to puncture (E p ) was calculated from the area under the force-displacement curve as a measure of efficiency. For unidirectional draw, only one prey item, H. plumieri (TL = 21.0-25.4 cm), was used. One side of the fish was adhered to a wood stage by cyanoacrylate glue. The stage was mounted into the MTS such that the mounted teeth were positioned with the 20

PAGE 32

cusp axis was parallel to the outward surface normal of the fish. The PVC coupler housing the teeth was in turn mounted to the 5 kN load cell. The cusp of each tooth was embedded in the epaxialis musculature (but not the pterygiophores) such that the entire cusp was embedded. Teeth sliced through the prey item at a velocity of 400 mm/s for 30 mm. Three trials were run for each tooth, but a new fish was used for each draw. For each trial, the maximum force that occurred during the draw event was recorded (F draw ). Following the draw trials, for a subsample of H. plumieri (n=11) the area surrounding the cut was skinned to determine the number of myosepta cut by the shark teeth. The number of cut myosepta was compared to the number of peaks in the force-displacement trace from the corresponding draw trial. Tooth morphometrics To capture tooth morphology, a series of tooth shape measurements was taken from five teeth from the functional row from five jaws of adult sharks for each species (Appendix A). Teeth chosen for measurements were from the same region of the jaw as those used in performance testing (Table 1.1). Only teeth from the right side of each jaw were used to account for any fluctuating asymmetry in tooth shape. The following measurements based in part on Shimada (2005) and on characters previously cited to be related to tooth function (Cappetta, 1987; Frazzetta, 1988; Abler, 1992; Motta, 2004) were taken (Fig. 1.2): base-cusp width (BCW; maximum cusp width), cusp height (CH; perpendicular from cusp apex to BCW), mesial cutting edge length (MCL; distance between cusp apex and most mesial point of BCW), and distal cutting edge (DCL; distance between cusp apex and most distal point of BCW). From these measurements, two ratios were 21

PAGE 33

calculated and used in subsequent analyses: cusp aspect ratio (CAR; CH/BCW) and cusp inclination (CI; MCL/DCL). In addition to the ratios, the following measurements were also used in the analyses: base overlap (BO; distance of either base overlap (+) or between bases (-) of adjacent tooth divided by the mean of BW for both bases), notch angle (NA; angle taken from cusp apex to notch to most distal point of BCW), and cusplet angle (CA; angle taken from cusp apex to notch to apex of lateral cusplets when present, CA = 0 if no cusplets present). Other aspects of tooth shape were quantified as discrete states: number of lateral cusplets (LAT), lingual-labial cusp curvature (LC; 0 = straight cusp, 1 = curved lingually, 2= recurved labially), and degree of serration (SE; 0 = none, 1 = weakly serrated (visible under a dissecting microscope), 2= strongly serrated (visible to the naked eye)). Statistical Analyses To determine if differences in puncture performance existed among tooth morphologies, means for force and energy values for each tooth within each prey item were taken. Means were compared using a two factor multivariate analysis of variance (MANOVA), with tooth and prey types as factors and individuals as replicates. Factors identified as significant were further assessed with two-way univariate analyses of variance (ANOVAs) and Holm-Sidak multiple comparison post-hoc tests. Analysis of interactions was not possible due to missing data from teeth that would not puncture, therefore to address differences within prey items and between teeth, one-way ANOVAs were performed for each prey item with Holm-Sidak multiple comparison or Tukeys HSD post-hoc tests. For each set of statistical tests, the following were log-transformed for 22

PAGE 34

normality: all puncture variables for H. plumieri, and F p and E p for C. sapidus. For those variables that either would not normalize or did not have equal variance (all puncture variables for E. saurus, F max for A. probatocephalus, S. tiburo and C. sapidus), non-parametric Kruskal-Wallis tests were run. Differences in draw performance (F draw ) among species were assessed using a one-way ANOVA and a post-hoc Tukeys HSD test. Paired t-tests were then used to determine whether tooth morphologies differed in puncture (F p F max ) and draw performance (F draw ). For all tests, a p-value of 0.05 was used to determine significance. MANOVA was performed using SYSTAT 11 (Systat Software Inc., San Jose, CA, USA); all other statistical tests were performed using SigmaStat 3.1 (Systat Software Inc., San Jose, CA, USA). Despite the fact that tooth morphology was not chosen based on taxonomy or evolutionary relationships, it is possible that the performance of different teeth is related to phylogeny. Simple exact Mantel tests were used to determine whether dissimilarity matrices of F p F max and E p for each prey item and F draw were correlated with a dissimilarity matrix based on phylogenetic distance. Distances were obtained from a composite tree based on the literature (Fig. 1.1) (Compagno, 1988; Martin et al., 1992; Naylor, 1992; Shirai, 1996; Grogan and Lund, 2004). Branch lengths were set to the first appearance of each extant species in the fossil record, though this underestimates divergence times, or to the age of the fossil tooth (first appearance data from Cappetta, 1987; Long, 1993; Cigala-Fulgosi, 1995; Iturralde-Vinent et al., 1996; Purdy, 1996; Yabe, 1998; Purdy et al., 2001; Rana et al., 2006). A significance level of p=0.05 was used for all tests. Mantel tests were performed with zt software (Bonnet and Van de Peer, 2002) (software available from http://www.psb.ugent.be/~erbon/mantel/). 23

PAGE 35

Lastly, canonical correlates analysis (CCA) was used to examine the relationship between tooth morphometric and performance variables. Due to problems with colinearity, two separate CCAs were run. The first encompassed performance variables from puncture testing on H. plumieri (F p F max and E p ) and morphometric variables CAR, CI, NA, CA, BO, LAT, LC, and SE. The second CCA used F draw and F p from H. plumieri and CAR, CI, NA, BO, and SE. CA, LAT, and LC all had the same values for the species that were capable of draw and therefore could not be included in the second CCA. CCAs were performed with SYSTAT 11 (Systat Software Inc., San Jose, CA, USA). All procedures were in accordance with University of South Florida Institutional Animal Care and Use Committee protocol number T2893. Results Puncture The majority of teeth successfully penetrated all prey items (Tables 1.2-1.4), and a representative force-displacement trace for a puncture may be found in Figure 1.3. Teeth that failed to penetrate were left out of subsequent statistical analyses. All puncture performance variables (F p F max and E p ) were significant for both tooth and prey factors (MANOVA, Wilks Lambda=0.061, p<0.001). Two-way ANOVAs on each of the puncture variables revealed differences in both tooth and prey factors for all variables (p<0.001) (Table 1.5). For both F p and F max the lowest forces were recorded during puncture of ladyfish E. saurus, while the highest forces occurred during puncture of blue crab C. sapidus (Table 1.2, 1.3, and 1.5). While both F p and F max were lower for the white grunt H. plumieri than for the sheepshead A. probatocephalus, neither were different 24

PAGE 36

from the bonnethead S. tiburo for F p Puncturing the S. tiburo only differed from A. probatocephalus for F max with lower magnitudes for S. tiburo (Tables 1.2 and 1.3). Elops saurus had lower E p than all prey items except H. plumieri, and only A. probatocephalus was greater than H. plumieri for E p (Table 1.4 and 1.5). No other differences in E p were found. Significant differences also occurred among teeth pooled across prey items, though no pattern emerged (Tables 1.2-1.5). F p for Cladodus sp., Xenacanthus compressus, and Carcharodon carcharias were greater than those of Negaprion brevirostris, Sphyrna mokarran, and Scymnodon ringens, though there were no differences within the groups. Cladodus sp. F p was also greater than that of both anterior and lateral Isurus oxyrinchus teeth. The same pattern exists for F max with the addition of Cladodus sp.and X. compressus also having higher magnitudes of force than Carcharhinus limbatus and Carcharhinus leucas, and X. compressus with greater F max than both I. oxyrinchus teeth. Other teeth only differed in magnitudes of F max and/or E p ; Prionace glauca, C. leucas, and Galecerdo cuvier had greater F max than S. ringens, N. brevirostris, and C. limbatus. For E p S. ringens was lower than C. leucas and G. cuvier, and G. cuvier was higher than C. limbatus. Within prey items, significant differences among teeth occurred, but there was no consistent pattern (Tables 1.2-1.4 and 1.6). No significant differences among teeth were found for any puncture variables for E. saurus or C. sapidus, nor were there differences in F p or F max within A. probatocephalus or S. tiburo. For the latter prey items, E p for Cladodus sp. was greater than that of C. carcharias, C.limbatus, N. brevirostris and both I. oxyrinchus teeth (Table 1.4 and 1.6). Additionally, for A. probatocephalus, E p for 25

PAGE 37

Cladodus sp. was greater than S. ringens, and C. leucas was greater than N. brevirostris. For S. tiburo, E p for Cladodus sp. was greater than C. leucas. For H. plumieri, all three puncture variables showed significant differences between C. carcharias and other teeth. For F p C. carcharias was higher than that of C. limbatus, Hybodus sp., and the anterior tooth of I. oxyrinchus. Carcharodon carcharias F max on H. plumieri was higher than that of Hybodus sp., both I. oxyrinchus teeth, and N. brevirostris. Lastly, E p for C. carcharias was greater than that of C. limbatus and the anterior tooth of I. oxyrinchus. Unidirectional draw For all teeth with successful draw tests, F draw was significantly higher than force at initial puncture F p (t 8 =-4.891, p=0.001), but was not different from F max of puncture (t 8 =0.821, p=0.433) (Table 1.7). Differences in F draw existed among tooth types (F 8,18 =2.258, p=0.018), however the only post-hoc difference found was that Cladodus sp. was higher than both G. cuvier and S. mokarran. A representative force-displacement trace may be found in Figure 1.4. Out of the 11 H. plumieri examined post-draw, the number of myosepta cut matched the number of peaks on the force-displacement trace for 54.5% of the specimens, 18.2% differed by one, and 27.3% did not match at all. Phylogenetic & morphometric relationships Performance is not related to phylogeny for the species tested in this study, as there was no apparent phylogenetic signal in the data. Mantel tests did not show any 26

PAGE 38

correlation between phylogenetic distance and any of the performance variables (p>0.05 in all cases). Likewise, there is little correlation between shape and performance. The first CCA (incorporating F p F max and E p ) indicated no correlation of puncture variables with shape or serrated versus non-serrated edges; no significant axes were produced by this analysis. The second CCA (incorporating only F draw and F p ) showed no relationship between tooth shape and draw performance. However, this CCA had one significant axis with F p SE, and NA loading positively on this axis, while CAR loaded negatively. As cusps became broader (CAR decreases), the degree of serration increased, notch angles became less acute, and the force to initial puncture increased. Discussion Performance provides the link between an organisms phenotype and its resource use (Wainwright, 1994; Koehl, 1996). Consequently, it is assumed that under adaptive scenarios there is predictive power between an animals morphology and its ecology or behavior (Weins and Rotenberry, 1980; Losos, 1990; Motta and Kotrshal, 1992). By investigating the biomechanical performance of shark teeth and seeking relationships to their feeding ecology, aspects of tooth shape that are predictive of tooth performance may be identified. Biomechanics of prey items Differences in puncturing performance occurred among different prey items. From a functional perspective, prey are often categorized into two groups: hard prey, 27

PAGE 39

such as echinoderms, crustaceans, and shelled mollusks, and soft prey (everything else) (Wainwright, 1988; Waller and Baranes, 1991; Turingan, 1994; Turingan et al., 1995; Dean et al., 2007; Ramsay and Wilga, 2007; Huber et al., in press). In terms of tooth biomechanics, this analysis questions this dichotomy, revealing that not all soft prey are alike. Even within a general prey type, such as teleosts, significant differences in puncture forces were found. Among teleosts, E. saurus required the lowest F p and F max while A. probatocephalus required the highest. Differences in efficiency (E p ) followed the same pattern; E. saurus required the least amount of energy for puncture, while A. probatocephalus required the most. This implies that not only does it generally take less force to puncture E. saurus, but less energy must be expended by the shark to do so. In addition, some teeth were able to penetrate some teleosts and not others. For example, teeth from the blue shark P. glauca could not puncture E. saurus or A. probatocephalus, but were able to penetrate H. plumieri, indicating that differences between prey items affect the function of some teeth. The thickness of the scales and skin, myoseptal arrangement, and muscle composition all likely play a role in how much force is required to puncture a particular teleost. Skin thickness has been shown to influence skin strength in teleosts (Hebrank and Hebrank, 1986) and penetration forces of knives into artificial skin (Gilchrist et al., 2008). Though not scales, one study found that forces produced by felid canine teeth puncturing the skin of pigs and deer were significantly different, even after the thick fur had been removed from the deer hide (Freeman and Lemen, 2007). Attachment of the scales to the body of the prey item may also be functionally important. For example, E. saurus has deciduous scales that are readily dislodged whereas A. probatocephalus scales 28

PAGE 40

are rarely dislodged. During puncture testing on E. saurus, large amounts of deformation occurred around the tooth, creating a crater-like depression and unpunctured scales were often found stuck to the tooth cusp. It is possible that as the sides of the crater become more vertical and the tooth continues to press on the edges of the scales, they may loosen and slip out of the way, creating a scaleless surface for the tooth to penetrate. Differences in skin architecture and stiffness in the bodies of prey items may contribute to the differing tooth puncture results both within and among the soft prey items. Skin tension is directly related to both the force and energy of knife penetration during stabbing (Gilchrist et al., 2008). Cross-helically wound collagen fibers contribute directly to skin stiffness in both teleosts and elasmobranchs; the number of collagen fiber layers varies among fishes, and even between sexes of some sharks (Motta, 1977; Hebrank, 1980; Hebrank and Hebrank, 1986; Pratt and Carrier, 2001). Similarly, strength and stiffness have been shown to be correlated with the amount and orientation of collagen fibers in mammalian skin (Flint et al., 1984; Vogel, 1988). Surprisingly, F p for the bonnethead shark S. tiburo were not different from either the white grunt H. plumieri or sheepshead A. probatocephalus, and F max for A. probatocephalus was greater than that of S. tiburo. In general, it has been assumed that shark skin is stiffer, as for a given body size sharks generally have thicker skin compared to teleosts (Hebrank and Hebrank, 1986). However, skin is not the only determinant of body stiffness. The perceived body stiffness is also a function of muscle, scales, and skeletal elements; differences exist between these factors for teleosts and sharks. For example, in sharks collagenous mysosepta are thickened as they attach to the dermis, but are not in most teleosts (Willemse, 1972; Hebrank and Hebrank, 1986). While the specific stiffness of these prey 29

PAGE 41

items are not known, it may be inferred from the results that one teleost was less stiff than other prey items; E. saurus was unique in that it would occasionally stay deformed following removal of shark teeth during puncture tests. This prey item required the least amount of force (F p and F max ) during puncture (Tables 1.2 and 1.3). Differences in force to puncture (F p ) among teleosts may explain why some shark species undergo positive allometric increases in bite force over ontogeny when no apparent ontogentic shift in diet occurs. For example, the blacktip shark Carcharhinus limbatus undergoes an allometric increase in bite force through ontogeny yet remains piscivorous during its entire life history (Dudley and Cliff, 1993; Castro, 1996; Barry, 2002; Hoffmayer and Parsons, 2003; Bethea et al., 2004; Huber et al., 2006). In addition to an increase in prey size throughout ontogeny, different teleosts occur in the diet of C. limbatus at different life stages (Bethea et al., 2004), possibly reflecting differing biomechanical challenges. Biomechanics of extant teeth Despite past efforts to classify sharks teeth into functionally relevant groups based on performance (Moss, 1977; Cappetta, 1987; Frazzetta, 1988; Motta, 2004), little support was found for biomechanical morphotypes. Differences in puncture performance occurred among tooth morphologies, both within prey items and when prey items were pooled, though few overall patterns occurred. Teeth of C. leucas, which are triangular and serrated, were less efficient during puncture (higher E p ) of A. probatocephalus than the more thinly cusped tooth of N. brevirostris. The broadly triangular Carcharodon carcharias teeth exhibited higher initial puncture forces (F p ) on H. plumieri than thin 30

PAGE 42

cusped C. limbatus and anterior I. oxyrinchus teeth, higher maximum puncture forces (F max ) than both I. oxyrinchus and N. brevirostris, and lower efficiency (E p ) than C. limbatus and I. oxyrinchus. This suggests that for some instances, it appears that broader triangular teeth, such as those of the white shark C. carcharias and the bull shark C. leucas are less efficient (smaller E p ) and require more force for puncturing compared to narrower teeth. These data are supported by the results of the CCA, though the white shark teeth being larger than the others might also account for higher puncture forces. Force at initial puncture (F p ) on H. plumieri increases as CAR (cusp aspect ratio) decreases; in other words, the broader and shorter the tooth cusp, the more force it takes to initially puncture H. plumieri. This result is consistent with other studies on cutting, for both mammalian teeth and knives (Freeman and Lemen, 2007; Gilchrist et al., 2008). However, broader teeth (as evidenced by a lower CAR) did not consistently perform more poorly at puncturing for all prey types. For the softest prey item, E. saurus, there was no difference in any performance variable among different extant teeth. Additionally, there was no difference in F max or F p for extant teeth puncturing A. probatocephalus or S. tiburo. In addition to contrasting patterns of puncture performance, there were surprisingly no differences in draw performance among tooth morphologies. Historically, tooth morphology has been the primary means of describing the biological role of teeth. In this regard, shark teeth in this study have been grouped in two general categories: tearing and cutting morphs (Cappetta 1987). Tearing teeth are described as having narrow cusps with distinct cutting edges, e.g. the anterior teeth of Isurus. Cutting teeth can be further divided into two subgroups: cutting subtype, where teeth are lingo31

PAGE 43

labially flattened, with broad cusps distally inclined toward the back of the jaw. Serrations are thought to further improve cutting performance. Teeth of Carcharodon carcharias, Galeocerdo cuvier, Hexanchus griseus, Carcharhinus leucas, Carcharhinus limbatus, Prionace glauca, Sphyrna mokarran, and the lateral teeth of Isurus oxyrinchus are members of this subgroup. The other subgroup is the cutting-clutching subtype. For this study, the C. limbatus and S. ringens teeth are part of the cutting jaw, while teeth of N. brevirostris are part of the clutching jaw. Differences in draw and puncture performance were not consistent with these groupings. Similarly, when comparing draw forces among extant teeth, no tooth was better or worse than another. Further, the canonical correlates analysis (CCA) showed that for successful draw tests, there was no correlation between tooth shape and force. Inclination, serration, and notching are cited as being characteristics of cutting teeth (Cappetta, 1987; Motta, 2004), yet our data did not support a performance benefit to this morphology. It is possible that a shape parameter that was not measured, such as cutting edge sharpness, could correlate with draw performance. Testing the teeth in draw for only one prey item may also have lead to a lack of differences. Many of these species prey on a variety of prey items of different sizes and taxonomic affinities (Compagno, 1984a, b; Cortes, 1999). As teeth encounter different materials within and among prey, such as muscle, bone, cartilage, or crustacean exoskeleton, they may perform differently as they did during puncture. This is supported by the force-displacement traces for draw, which has several peaks (Fig 1.4). The number of myosepta cut matched the number of peaks on the force-displacement trace for the majority (54.5%) of the specimens, indicating that it may take more force to cut through a tough collagenous myoseptum than through muscle. 32

PAGE 44

As it has been suggested that myosepta not only transmit muscle forces, but also potentially constrain myomere bulging during contraction, these tissues should be relatively stiff (Gembella et al., 2003; Brainard and Azizi, 2005). When considering both biological roles of puncturing and cutting, the majority of teeth did not perform better in puncture (F p F max ) or unidirectional draw. Many extant teeth were able to both puncture and cut H. plumieri (Figure 1.1). Further, there were no differences in any performance variables for carcharhinid upper jaw or lower jaw teeth of the cutting-clutching subtype (C. limbatus and N. brevirostris, respectively). When taking large prey, some carcharhinid sharks will bite, which engages the teeth in puncture, then laterally head shake, which engages the teeth in draw (Frazzetta and Prange, 1987). In this case, teeth must perform both roles within the same predation event. Teeth that perform multiple functions have been found in other elasmobranchs as well. For example, the bamboo shark Chiloscyllium plagiosum has small clutching teeth, but will take both hard and soft prey. It is able to do so because the teeth passively rotate and flatten to form a flat surface for crushing (Ramsay and Wilga, 2007). The homodont teeth of the lesser electric ray Narcine brasiliensis also have dual roles; teeth on the occlusal surface are used for grasping, while teeth on the external surface may protect the jaws during excavation of prey (Dean et al., 2008). Additionally, teeth in elasmobranchs are used for gripping females during mating, and this has also been shown to be related to tooth morphology (Springer, 1967; Compagno, 1970; Fedducia and Slaughter, 1974; McCourt and Kerstitch, 1980; Compagno, 1988; Kajiura and Tricas, 1996). When structures have multiple functions, they may not be optimized for any one function, 33

PAGE 45

making correlations between form and function difficult (Reif, 1983; Lauder, 1995; Koehl, 1996; Domenici and Blake, 2000). For all teeth with successful draw tests, it took less force to make an initial puncture than to draw a tooth through the prey item, though the maximum force produced during puncture was equal to the force produced during draw. As a tooth continues to enter the prey item after initial puncture, the initial rift is widened while new tissue is being punctured at the tooth apex. One study found that this difference is likely only important for the initial puncture; for mammalian prey muscle and fat offered little resistance, implying that differences between sharp and blunt cusps are erased as most of the force generated during the remainder of the puncure is generated as the teeth widen the hole made by penetration (Freeman and Lemen, 2007). During initial puncture on teleosts prey items, only scales and skin are penetrated, but as the tooth continues through the prey item, it also encounters muscle tissue and myosepta. Since the draw experiments were carried out with the cusp already embedded in the prey item, it was encountering four materials throughout the entire test. Despite a general lack of specialization, it is clear that some of the teeth that were tested in both unidirectional draw and puncture performed better at one task or the other (Fig. 1.1). Scymnodon ringens could not cut during draw, but was able to puncture H. plumieri. This small serrated tooth is classified as cutting type (Cappetta, 1987). Likewise, the multicusped Hexanchus griseus teeth would not puncture H. plumieri, and the notched and heavily serrated Galeocerdo cuvier teeth had only one successful puncture, yet both teeth were able to cut during unidirectional draw. Tiger sharks, G. cuvier, routinely cut through large sea turtle shells by vigorously biting and shaking their 34

PAGE 46

head and teeth back and forth across the prey, employing draw (Heithaus, 2001; Simpfendorfer et al., 2001). If this comparison between puncture and draw tests is extended to other prey items, Prionace glauca could not puncture any prey except H. plumieri, while Sphyrna mokarran could not puncture A. probatocephalus or S. tiburo, yet both teeth could cut during draw. When all teeth that fail at puncturing are considered, a morphological pattern emerges. The cusp apices of these teeth are distally inclined to the point where the flat surface of the mesial cutting edge contacts the prey during puncture, not the tip of the tooth. As tooth tips become sharper, less surface area is touching the tooth. For a given force, this increases the amount of stress applied to the prey item, as stress is equal to force divided by area. Compressing the prey item during the initial stages of puncture creates compressive forces beneath the tooth tip and tension on the surrounding substrate as the prey item bulges around the tooth. These stresses together generate an effective shear stress which, if large enough, ruptures the tissues (Frazzetta, 1988). Increasing the amount of surface area of the tooth-prey contact decreases the amount of stress produced during puncture. The flat surface of these teeth may decrease stress on the prey item to the extent that puncture (failure) is not possible. This is further supported by studies on mammalian tooth shape, where blunter teeth required significantly more force to puncture insects (Evans and Sanson, 1998), fruits (Freeman and Weins, 1997) and mammalian hides (Freeman and Lemen, 2007), compared to sharp teeth. For sharks, however, the tension in the bulging tissue may facilitate cutting during unidirectional draw. As the tooth is moved distally during draw, the tip of the cusp can engage the tissue, creating compression directly under the cusp tip and further adding tension via more bulging (Frazzetta, 1988). 35

PAGE 47

There are other aspects of teeth that may contribute to tooth performance that need to be addressed. Teeth of the upper and lower jaws may also interact with each other. How teeth of the upper and lower jaws shear past each other and how teeth of the same jaw affect puncture and cutting during draw are unanswered questions. A study using utility blades as proxies for upper and lower jaw teeth indicated that certain combinations of blade shapes affect cutting efficiency (Anderson and LaBarbera, 2008). The flexible collagenous attachment of shark teeth may also facilitate cutting during draw as the teeth may pivot anteroposteriorly around obstructions preventing them from hanging-up on tough material (Frazzetta, 1988). In reality, sharks teeth form a functional complex that work together. Tooth base overlap within the same jaw may transmit forces to linked teeth, as overlapping bases are lashed together with collagenous Sharpeys fibers (Frazzetta, 1988). Including these parameters in future studies may further elucidate the link between tooth morphology and performance. Fossil tooth mechanics In general, fossil teeth did not perform well at unidirectional draw, but performed similarly to modern teeth in puncture. Out of the three fossil morphologies tested, Cladodus sp. was the only tooth that was successful at draw. Failure of the Hybodus sp. and X. compressus teeth to successfully draw may be due in part to a lack of sharpness during our tooth reconstruction. However, other aspects of morphology may have resulted in the lack of draw performance. Extant shark teeth with distally inclined cusps do not have the entire cusp inclined; instead only the apical portion of the cusp is inclined, forming a notch on the distal cutting edge. The distal cusp of X. compressus has 36

PAGE 48

no such notch as the entire cusp is distally inclined. Even teeth that are marginally inclined, such as the anterior tooth of I. oxyrinchus (CI = 1.12) and C. carcharias (CI = 1.05) have either a slight notch (NA for I. oxyrinchus = 154.02 degrees) or serrations, as in C. carcharias, where stress can be concentrated between serrations (Frazzetta, 1988; Motta, 2004). In contrast, Hybodus teeth have a functional notch between the main cusp and the first distal cusplet, though it is not bladed. All three fossil morphologies were successful at puncture, though to varying degrees. Cladodus sp. was able to puncture all prey items. Overall, Cladodus sp. produced higher forces (F p F max ) and was less efficient (E p ) than the majority of the extant teeth. Yet for the softest and hardest prey items (E. saurus and C. sapidus, respectively), the Cladodus tooth performed similarly to extant teeth. Considering all the performance tests, Cladodus teeth appear to function similarly to many of the extant teeth. This is surprising, as the majority of the concurrent possible prey items during the Paleozoic were armored, including placoderms, cephalopods, and osteichthyans with rhomboid scales (Brett and Walker, 2002). It is important to note that some soft prey were also available in the form of acanthodian fishes, conodonts, and other chondrichthyans. Direct and indirect evidence, in the form of preserved gut contents and trace fossils on potential prey items, exists for cladodont shark predation on both hard and soft prey (Mapes and Hansen, 1984; Williams, 1990; Mapes et al., 1995), and their teeth appear biomechanically suited for a variety of functional prey types. Xenacanthus compressus teeth were able to puncture E. saurus, H. plumieri, and C. sapidus, but not A. probatocephalus or S. tiburo. Overall, X. compressus produced higher forces (F p F max ) than the majority of the extant teeth and were less efficient at 37

PAGE 49

puncture than several modern teeth, but like Cladodus sp. were not different from extant teeth in puncturing E. saurus or C. sapidus. It is therefore possible that X. compressus was similarly capable of handling a variety of prey items, though perhaps not as diverse of a list as Cladodus. There is less dietary evidence for freshwater xenacanthid sharks than those marine sharks with cladodont-type teeth. One Permian xenacanthid, Triodus sessilis, has been described with two species of larval temnospondyl amphibians in the gut (Kriwet et al., 2008). Another study describes evidence that Orthacanthus, an Upper Carboniferous xenacanthid, fed on the xenacanthid Triodus (Soler-Gijon, 1995). While there is no evidence for durophagy in xenacanthid sharks, the teeth appear biomechanically suited for puncture of hard prey. The multicusped Hybodus sp. tooth was the least successful at puncturing soft prey items; only one trial on H. plumieri was successful. However, these teeth were able to puncture the blue crab C. sapidus. Cappetta (1987) designates the majority of Hybodus teeth as clutching-type (those Hybodus teeth with smaller central cusps), with others falling into tearing-type (those with a high central cusp). There is no dietary data for hybodont sharks, however one paleoecological study of trophic levels in Jurassic sediments proposed that Hybodus preyed upon fishes and squid, as well as scavenged, based on tooth shape and available prey items (Martill et al., 1994). The Hybodus tooth used in this study has a relatively small central cusp compared to the lateral cusplets, and the lack of soft prey puncture and draw supports a non-tearing role. As this tooth was only successful at puncturing hard prey, it appears suited for durophagy. 38

PAGE 50

Conclusions In this study, I investigated the performance of three general categories of extant teeth, tearing-type, cutting-type, and cutting-clutching type, as well as three fossil morphologies, on a variety of prey items. The goals of this study were to: (1) Determine the forces necessary for individual teeth to penetrate a variety of fish and crustacean prey representative of shark diets; (2) Determine what differences in penetration force and efficiency occur among tooth types; (3) Compare performance between different cutting regimes for a given tooth morphology; and (4) Determine which morphological aspects, if any, of tooth shape are predictive of tooth performance. Differences in puncturing performance occurred among different prey items, indicating that not all soft prey items are alike. The majority of teeth were able to puncture different prey items, and differences in puncture performance also occurred among tooth types; however, few patterns emerged. Force to puncture was less than the maximum force that occurred during draw tests, however there were no differences between the maximum draw forces and maximum puncture forces. Few morphological patterns were identified. In some cases, broader triangular teeth were less effective at puncturing than narrow-cusped teeth. Teeth from Galeocerdo cuvier, Prionace glauca, Hexanchus griseus, and Sphyrna mokarran were unable to puncture many of the soft prey items. The flat surface of the tooth-prey contact may decrease stress on the prey item to the extent that puncture (failure) is not possible. No morphological characteristics were correlated with maximum draw force. Many of the extant shark teeth in this study were not only able to perform draw and puncture equally well, but tooth morphologies were functionally equivalent to each other. This does not support the use of tooth morphology to predict biological role. 39

PAGE 51

Additionally, fossil morphologies were generally successful at puncture and overall similar in performance to extant teeth, but only Cladodus sp. was able to perform in draw. Clearly, we have just scratched the surface on shark tooth performance and it is difficult to conjecture about the evolution of sharks without more information. This study addressed the performance of isolated teeth, the majority of which are bladed. There are other morphologies of extant shark teeth, such as molariform teeth, the multicusped teeth of the prickly shark Echinorhinus cookei, and the unique tricuspid teeth of the frilled shark Chlamydoselachus anguineus, for which there is no performance data. While I have presented the first performance tests on fossil teeth, I have only sampled three teeth from over 400 million years of evolution. Even within broad generalizations of fossil shark tooth morphology (e.g. cladodont, xenacanth, hybodont), several different morphologies exist (Zangerl, 1981; Cappetta, 1987). Additionally, there are other aspects of teeth that may contribute to tooth performance that need to be addressed, including flexible tooth attachments (but see Ramsay and Wilga, 2007), tooth base overlap that may transmit forces to linked teeth (Frazzetta, 1988), and lateral cusplets. In reality, sharks teeth form a functional complex that works together. How teeth of the upper and lower jaws shear past each other and how teeth of the same jaw affect puncture and cutting during draw are unanswered questions. Including these parameters may elucidate the link between tooth morphology and performance. 40

PAGE 52

Figure 1.1: Phylogenetic tree of species included in this study and tooth morphology, based on Compagno, 1988; Martin et al., 1992; Naylor, 1992; Shirai, 1996; Grogan and Lund, 2004. Branches are not drawn to scale. a = anterior, l = lateral. Line drawings show tooth used for each species, shaded for morphotype: light grey = tearing, dark grey = cutting-clutching subtype, white = cutting subtype. H = able to penetrate hard prey, vertical arrow = able to puncture more than half of the soft prey items, horizontal arrow = able to perform unidirectional draw. 41

PAGE 53

Figure 1.2: Tooth measurements taken for canonical correlation analysis. BCW = base-cusp width, BO = base overlap, BW = base width, CA = cusp angle, CH = cusp height, DCL = distal cutting edge, MCL = mesial cutting edge, NA = notch angle. 42

PAGE 54

Figure 1.3: Force-displacement trace for puncture of Haemulon plumieri by an anterior Isurus oxyrinchus tooth. Arrow denotes initial puncture (F p ). 43

PAGE 55

Figure 1.4: Force-displacement trace for unidirectional draw through Haemulon plumieri by an anterior Isurus oxyrinchus tooth. Arrows indicate probable cuts through myosepta. 44

PAGE 56

Table 1.1: Tooth and jaw position information for each extant species in this study. N = number of individuals for each species. Position = tooth family counting from the jaw symphysis. Specific jaw position for C. carcharias is not known. Species nJaw Position I. oxyrinchus (a) 3Lower 2 I. oxyrinchus (l) 3Upper 6 C. carcharias 3Upper "Anterior" S. mokarran 3Upper 6 G. cuvier 3Lower 3 C. leucas 3Upper 8 C. limbatus 3Upper 5 P. glauca 3Upper 7 N. brevirostris 3Lower 3 S. ringens 1Lower 1 H. griseus 1Lower 2 45

PAGE 57

Table 1.2: Means for force to puncture (F p ) (N = Newtons) + standard error. n = number of teeth used for puncture tests on each prey item. X = puncture was not successful. Deviations from n given in Table 1 occurred when some teeth could not successfully puncture a given prey item, but others could. Species mean = mean for each species with prey items pooled. Prey mean = mean for each prey item with species pooled. SpeciI. ox esE. saurusH. plumieriA. probatoc.S. tiburoC. sapidusSpecies Meanyrinchus (a)7.36 + 1.945.66 + 1.8518.49 + 6.6111.38 + 1.9335.23 + 4.0715.93 + 3.42yrinchus (l)5.00 + I. ox 2.728.31 + 2.6421.05 + 5.4011.08 + 2.2733.20 + 10.5515.98 + 3.88 carcharias15.85 + C. 2.8022.66 + 0.8034.25 + 2.7016.68 + 1.4465.12 + 15.4031.03 + 5.57. mokarran1.125.57 + S 0.23XX23.29 + 4.2413.69 + 4.74. cuvierXX45.54X70.52 + G 21.7264.28 + 16.58 leucas4.95 + C. 1.149.82 + 2.1627.00 + 9.3123.0551.76 + 5.1424.7 + 6.04. limbatus1.27 + C 0.164.85 + 1.017.84 + 1.585.1938.13 + 7.4213.85 + 5.06 glaucaX4.06XXX4.055. brevirostris3.78 + P.N 2.705.05 + 2.3710.61 + 8.8013.62 + 4.1127.98 + 5.2912.98 + 3.12ngens0.3821.444.33XX8.72 + S. ri 6.46 griseus1.98XXX89.4145.70 + H. 43.72odus sp.X6.30XX68.4037.35 + Hyb 31.05 compressus10.1950.24XX62.0040.81 + X. 15.68ladodus sp.15.7712.2666.75112.7779.0357.32 + C 19.23rey Mean6.70 + P 1.3210.98 + 2.1023.99 + 4.1621.43 + 7.7647.66 + 4.60 46

PAGE 58

Table 1.3: Means for maximum force during puncture (F max ) (N = Newtons) + standard error. n = number of teeth used for puncture tests on each prey item. X = puncture was not successful. Deviations from n given in Table 1 occurred when some teeth could not successfully puncture a given prey item, but others could. Species mean = mean for each species with prey items pooled. Prey mean = mean for each prey item with species pooled. SI. ox peciesE. saurusH. plumieriA. probatoc.S. tiburoC. sapidusSpecies Meanyrinchus (a)9.49 + 2.2815.73 + 0.4526.46 + 2.1413.54 + 0.2383.03 + 22.0530.80 + 3.42yrinchus (l)8.64 + I. ox 3.2711.46 + 2.2824.56 + 4.4915.178 + 1.11105.63 + 10.9435.13 + 11.48archarias22.13 + C. c 0.3739.50 + 2.7540.55 + 6.4721.29 + 2.07158.91 + 12.0256.48 + 14.07okarran7.7015.59 + S. m 1.23XX94.67 + 5.9153.81 + 18.50X38.9552.25X149.58 + G. cuvier 21.92107.99 + 28.24 leucas8.15 + C. 0.4123.95 + 4.8540.93 + 4.3122.95142.23 + 32.7456.33 + 18.59 limbatus5.14 + C. 0.1812.38 + 2.8117.08 + 1.305.7892.66 + 13.6433.21 + 11.75X53.37XXX53.37virostris5.89 + P. glaucaN. bre 1.177.92 + 2.0221.28 + 6.1413.22 + 3.4066.41 + 12.7124.32 + 7.32ens5.748.318.88XX7.64 + S. ring 0.97us5.85XXX198.69102.27 + H. grise 96.42X17.36XX118.9368.15 + Hybodus sp. 50.79ompressus26.30107.92XX208.58114.27 + X. c 52.7218.6132.1085.42135.22119.9878.27 + Cladodus sp. 23.16e Pr y Mean11.13 + 1.6223.92 + 4.1932.66 + 4.4924.99 + 9.31118.77 + 8.57 47

PAGE 59

Table 1.4: Means for energy to puncture (E p ) (N/mm) + standard error. Larger numbers indicate less efficiency. n = number of teeth used for puncture tests on each prey item. X = puncture was not successful. Deviations from n given in Table 1 occurred when some teeth could not successfully puncture a given prey item, but others could. Species mean = mean for each species with prey items pooled. Prey mean = mean for each prey item with species pooled. SpeI. ox ciesE. saurusH. plumieriA. probatoc.S. tiburoC. sapidusMeanyrinchus (a)0.028 + 0.0090.017 + 0.0040.059 + 0.0250.031 + 0.0070.019 + 0.0060.028 + 0.007yrinchus (l)0.011 + I. ox 0.0070.018 + 0.0060.061 + 0.0220.024 + 0.0090.018 + 0.0100.024 + 0.006. carcharias0.071 + C 0.0270.155 + 0.0430.116 + 0.0280.061 + 0.0100.054 + 0.0200.091 + 0.014okarran0.0020.013 + S. m 0.005XX0.008 + 0.0010.008 + 0.002. cuvierXX0.142X0.077 + G 0.0230.086 + 0.025eucas0.013 + C. l 0.0020.019 + 0.0050.165 + 0.0580.0550.031 + 0.0080.051 + 0.019batus0.002 + C. lim 0.0000.010 + 0.0030.017 + 0.0010.0070.024 + 0.0080.013 + 0.003aucaX0.006XXX0.006. brevirostris0.009 + P. glN 0.0070.010 + 0.0060.003 + 0.0030.019 + 0.0090.023 + 0.0100.014 + 0.0040.00020.0030.008XX0.004 + S. ringens 0.002 griseus0.003XXX0.0450.024 + H. 0.021odus sp.X0.009XX0.0520.030 + Hyb 0.021pressus0.0330.153XX0.0390.075 + X. com 0.039odus sp.0.0930.0250.3050.810.0790.262 + Clad 0.145n0.025 + Mea 0.0070.036 + 0.0120.087 + 0.0200.094 + 0.060.034 + 0.005 48

PAGE 60

Table 1.5: Two-way ANOVA results for pooled species and pooled prey. F p = force at initial puncture, F max = maximum puncture force, E p = Energy to puncture, df = degrees of freedom. The error degrees of freedom are 83 for all tests. = p<0.05. FpFmaxEpdf131313F5.8414.459.56p<0.001*<0.001*<0.001*df444F37.71139.799.03p<0.001*<0.001*<0.001*SpeciesPrey 49

PAGE 61

Table 1.6: One-way ANOVA results for species within prey items. F p = force at initial puncture, F max = maximum puncture force, E p = Energy to puncture, df = degrees of freedom. # = non-parametric Kruskal-Wallis test performed instead of ANOVA, = p<0.05, + = no differences found in post-hoc tests, therefore results are considered statistically nonsignificant. FpFmaxEpdf101010F14.93#15.28#16.21#p0.140.120.09df6, 136,136,13F6.825.264.46p0.002*0.006*0.011*df5,885,8F1.7614.11#5.04p0.280.080.022*df63,63,6F8.54#2.464.88p0.200.160.048*df7,16117,16F3.1318.46#2.81p0.03+0.070.04+C. sapidusE. saurusH. plumieriA. probatoc.S. tiburo 50

PAGE 62

Table 1.7: Means for maximum force during draw (F draw ) (N = Newtons) + standard error. n = number of teeth used for puncture tests on each prey item. X = draw was not successful. SpeciesFdrawI. oxyrinchus (a)28.33 + 5.48I. oxyrinchus (l)13.90 + 1.63C. carcharias26.53 + 2.96S. mokarran13.58 + 3.78G. cuvier12.14 + 3.6C. leucas17.14 + 2.82C. limbatus15.02 + 2.47P. glauca15.96 + 2.40N. brevirostris21.24 + 2.48S. ringensXH. griseus17.55Hybodus sp.XX. compressusXCladodus sp.37.39 51

PAGE 63

52 Chapter 2: Biology Meets Engineering: Th e Structural Mechan ics of Shark Teeth Abstract Teeth are an integral part of the vertebrate feeding apparatus. They can divide prey into manageable pieces, grip prey for reorientation, and process prey to remove inedible components. Despite this, the majority of st udies on the evolution and function of feeding in sharks have focused primarily on the movement of cranial components and muscle function, with little integrati on of tooth properties or f unction. Those biomechanical studies that have addressed elasmobranch tooth form and function have largely focused on the qualitative mechanics of cutting, ignorin g the biomechanics of the tooth itself. As teeth are subjected to sometimes extreme loads during feeding, they undergo stress, strain, and potentially failure. While performa nce is certainly subject to natural selection, attributes related to structur al strength such as material properties and overall shape may also be subjected to natural selection. Therefore, both prey processing ability and structural parameters must be considered to understand the evoluti on of shark teeth. In this study, the structural mechanics of fossil and extant shark teeth are investigated. The first goal was to determine the material properties for enameloid, osteodentine, and orthodentine via nanoindentation. While shark de ntines are harder than other vertebrate dentines, enameloid has similar hardness and Youngs modulus to mammalian enamel. This latter relationship may be due to sim ilar microstructures between shark enameloid

PAGE 64

53 and mammalian enamel. I then used finite element analysis to visualize stress distributions of fossil and extant shark teeth during puncture, unidirectional draw (cutting), and holding. The specific goals for th is portion of the study were to determine if tooth morphologies are more structurally strong during diffe rent loading regimes and to examine the role of morphological features, su ch as notches or cusp shape, on stress distribution. Under the loading and bounda ry conditions in this study, which are consistent with bite forces of large shar ks, shark teeth are structurally strong. Teeth loaded in puncture have localized stress concentrations at the cusp apex that diminish rapidly away from the apex. When loaded in draw and holding, the ma jority of the teeth show stress concentrations cons istent with mechanically sound cantilever beams. Notches result in stress concentration during draw a nd may serve as a weak point; however they are functionally important for cutting prey during lateral head shaking behavior. As shark teeth are replaced regularly, it is proposed that the frequency of tooth replacement in sharks is driven by tooth wear, not tooth failure. Introduction Teeth are an integral part of the vertebrate feeding apparatus. They can divide prey into manageable pieces, grip prey for reorientation, and process prey to remove inedible components. Despite this, the majority of st udies on the evolution and function of feeding in sharks have focused primarily on the movement of cranial components and muscle function, with little integration of tooth properties or functi on (Motta, 2004; Huber et al., 2005; Dean et al., 2007; Ramsay and Wilga, 2007; Dean et al., 2008; Motta et al., 2008). Continuous tooth replacement, coupled with the cartilaginous nature of the skeleton, has

PAGE 65

54 led to a chondrichthyan fossil record composed primarily of teeth. To understand the evolution of the shark feeding mechanism, we must understand the contribution of all parts of the feeding apparatu s, including the teeth. Extant shark teeth encompass a wide va riety of shapes, including teeth with triangular serrated cusps, oblique serrated a nd non-serrated cusps, not ched serrated cusps, non-serrated recurved cusps, multicusped teet h, and flattened tooth pavements. The forms are often ascribed qualitativ e functions without any biom echanical testing, employing terminology such as gripping, piercing, cr ushing, cutting, or tearing (Cappetta, 1986, 1987; Motta, 2004). Biomechanical studies that have addresse d elasmobranch tooth form and function have largely focused on the qualitative mechanic s of cutting itself (Frazzetta, 1988; Abler, 1992). More recent st udies focus on reorientation of teeth during feeding and their dual roles in gripping and either crushing or protecting the jaw as it hits the seafloor (battering) (Ramsay and Wilga, 2007; Dean et al., 2008). In general, the biomechanics of the tooth itself have been ignored. While the forces necessary for teeth to penetrate teleost and elasmobranch prey items are on the order of tens of Newtons (s ee Chapter 1 of this dissertation), some chondrichthyans are capable of producing far hi gher bite forces. Some elasmobranchs fall on the low end of the scale; static equilibrium models calculate the average anterior bite force for Etmopterus spinax to be 1 N (mean SL = 32.5 cm) (Claes & Malefet, unpublished data), while a bite force of 20 N was calculated for a 45.1 cm SL Squalus acanthias (Huber and Motta, 2004). Many chondricthyans fall between 100 and 1000 N of bite force. The spotted ratfish Hydrolagus colliei (44 cm TL) produced 58 N of anterior bite force under teta nic stimulation, and static e quilibrium models estimated

PAGE 66

55 anterior bite force of 104 N and 191 N pos teriorly (Huber et al., 2008). The horn shark Heterodontus francisci (74 cm TL) produced a restrained anterior b ite force of 187 N, but based on static equilibrium models is capable of 128 N anteriorly and 338 N posteriorly (Huber et al., 2005). Static e quilibrium models estimated th e bite force of a 152 cm TL Carcharhinus limbatus to be 423 N anteriorly and 1083 N posteriorly (Huber et al., 2006) and a mean anterior bite force of 834 N for the bull shark Carcharhinus leucas (170 cm mean SL) (Huber & Mara, unpublished da ta). The highest calculated bite forces are far greater than 1000 Newtons Static equilibrium models estimate the posterior theoretical bite force of the great hammerhead Sphyrna mokarran (4.3 m TL) to be 6080 N (Mara et al, unpublished data); finite element models of a 6.4 m TL white shark, Carcharodon carcharias estimate bite forces of 9320 N anteriorly and 18216 N posteriorly (Wroe et al., 2008). As teeth are subjected to these sometimes extreme loads during feeding, they undergo stress, strain, and potentially fail ure. In order to continue performing their biological role, teeth must resist breakage until they are shed. It is unclear whether tooth morphologies are more structurally strong under one loading regime versus another. For example, do cu tting teeth fail less often during cutting compared to puncture? While performance is certainly subject to natural selection (Arnold, 1983; Bennet, 1991), attrib utes related to structural strength such as material properties and overall shape may also be subj ected to natural select ion (Erickson et al., 2002; Lucas et al., 2008). Therefore, both prey processing ability and structural parameters must be considered to unde rstand the evolution of shark teeth.

PAGE 67

56 Shark tooth materials Shark teeth are composite material structures. Each tooth can be divi ded into two zones: the crown and the root or base. The tooth can be further divide d into two distinct structural components: a centr al core of dentine covered by enameloid, an enamel-like substance formed from both odontoblasts and ameloblasts (Poole, 1967a). Some elasmobranchs also have a central pulp cavit y (orthodont teeth), while in others the root extends into the central core of the crown (osteodont teet h) (Cappetta, 1987; Compagno, 1988). Enameloid is a highly mineralized tissue, composed primarily of hydroxyapatite crystallites. These crystallites are arranged in bundles that vary in orientation depending on the location within the tooth (Poole, 1967a; Gillis and Donoghue, 2007). In modern sharks, these bundles tend to align into three layers: a thin su perficial layer with randomly arranged single cystallites (s hiny layer enameloid, SLE), followed by a parallel-fibered layer (PFE), and an inne rmost tangled-fiber layer (TFE) (Gillis and Donoghue, 2007). Previous work has shown that the PFE imparts tens ile strength, while the TFE provides resistance to compressive forces (Preuschoft et al., 1974). Together, shark enameloid forms a layer that is 0.2-0.9 mm thick (Preus choft et al., 1974). Elasmobranch dentine is composed of mineralized collagenous tissue (Bradford, 1967; Johansen, 1967). Two types of dentine oc cur. Osteodentine superficially resembles spongy bone; the dentine surrounds vascular canal s, similar to osteons. Orthodentine does not contain dental osteons. In stead it contains smaller para llel branching tubules that provide a banded appearance (Compagno, 1988). Os teodentine forms the base in all shark teeth. The crown can be composed primarily of either orthodentine (orthodont), such as

PAGE 68

57 the teeth of carcharhiniform sharks, or osteode ntine (osteodont), as in lamniform sharks (Mertinene, 1982; Compagno, 1988). Despite a prolific data base on mammalia n tooth properties, especially humans (Waters, 1980; Brear et al., 1990) there is a paucity of data on fish tooth material properties. While the material properties ha ve been determined for elasmobranch jaw cartilage (Summers and Long, 2006) and verteb rae (Porter et al., 2006; Porter et al., 2007), the material properties of shark teeth and their components have not been studied to date. In this study, the structural mechanic s of fossil and extant shark teeth are investigated. The first goal was to determine the material properties for enameloid, osteodentine, and orthodentine via nanoindent ation. The hardness and elastic modulus of these tissues is not known, and is necessary for the second portion of this study. I then used finite element analysis to visualize st ress distributions of fossil and extant shark teeth during puncture, unidirect ional draw (cutting), and holding. The specific goals for this portion of the study were to determine if tooth morphol ogies are more structurally strong during different loading regimes a nd to examine the role of morphological features, such as notches or cusp shape, on stress distribution. Materials & Methods Nanoindentation Five teeth each of one carchar hiniform species, the bonnethead Sphyrna tiburo, and one lamniform, the sand tiger shark Carcharias taurus, were utilized to compare teeth with orthodont and osteodont dentine. Freshly shed teeth from C. taurus were collected from

PAGE 69

the floor of the Living Seas aquarium exhibit (Epcot Center, Walt Disney World, Florida, USA). Anterior teeth from S. tiburo were obtained from a 34.5 cm TL female that was euthanatized approximately one hour before teeth were collected. Teeth from both species were stored in seawater. Prior to testing, teeth were removed from the seawater and dried completely. Each tooth was sectioned transversally (relative to the tooth) approximately halfway down the cusp to create the surface for indentation, then again at the base to create a flat surface for mounting. Testing surfaces were polished with 400 grit sandpaper for four minutes and finally with Pikal polishing paste (Nihon Maryo-Kogyo Co., Japan) for twenty minutes to ensure a smooth surface. The polished samples were then washed in ethanol to remove any debris, and mounted on stainless steel cylinders with cyanoacrylate glue. Specimens were tested with a MTS Nanoindenter XP (Eden Prairie, MN, USA) with a Berkovich diamond tip. Both enameloid and dentine were tested on each tooth (Figure 2.1), with nine 2 micron deep indentations preformed on each material. Indentation sites were chosen haphazardly. Specimens were indented with a target strain rate of 0.5 s -1 Load and displacement were continuously recorded throughout the indentation process for each tooth and tooth material. For each indentation, the resulting load-displacement curve was used to calculate hardness (H) and Youngs modulus (E). Hardness was calculated at the peak load, and is given by Eq. (1): A WHmax (1) 58

PAGE 70

where W max is the peak load (N) and A is the projected indentation area (M 2 ) of the Berkovich indenter (Fisher-Cripps, 2004). The reduced Youngs modulus for the contact (Er), that is of the indenter and the specimen, was determined from the slope of the unloading curve at the maximum load, and is given by Eq. (2): dhdWAEr21 (2) where dW/dh is the slope of the unloading curve (Fisher-Cripps, 2004). The reduced Youngs modulus is related to the Youngs modulus of the specimen being indented (E s ) by Eq. (3): ssiir E v E v E )1()1(122 (3) where v i is the Poissons ratio of the indenter, E i is the Youngs modulus of the indenter, and v s is the Poissons ratio of the specimen being indented. As Poissons ratio is unknown for shark enameloid or dentine, the Poissons ratio for mammal enamel and dentine were used (v = 0.3 for both materials) (Waters, 1980). Many materials, including biological materials, have Poissons values of 0.3, so this is not an unreasonable assumption (Vogel, 2003; Wroe et al., 2008). All calculations of hardness and Youngs modulus were done in TestWorks 4 (MTS Systems Corporation, Eden Prairie, MN, 59

PAGE 71

60 USA). Means of hardness and Youngs modulus for each material were taken for each tooth. T-tests were then used to determine if there were differences in hardness and Youngs modulus between the tooth materials of the two species ( p=0.05) with SigmaStat 3.1 (Systat Software Inc., San Jose, CA, USA). Finite Element Analysis Specimens Teeth from ten shark species were chosen to c over a wide range of extant tooth forms, as opposed to taxonomy: Carcharhinus leucas, Carcharhinus limbatus, Carcharodon carcharias, Galeocerdo cuvier, Hexanchus griseus, Isurus oxyrinchus, Negaprion brevirostris, Prionace glauca, Scymnodon ringens, and Sphyrna mokarran (Table 2.1, Figure 2.2). Note that two t eeth (one lateral and one an terior) were chosen from Isurus oxyrinchus due to the presence of diversity of tooth shapes within the jaw. We also chose a single tooth from three fossil species to re present basic fossil morphologies that are not found in modern sharks: cladodont ( Cladodus sp. Stephens Museum 1998-1a), xenacanth ( Xenacanthus compressus, USNM 182325), and hybodont ( Hybodus sp. USNM 14197). Modeling The geometry of each tooth was acquired via a Phillips Mx8000 high-resolution x-ray computed tomography scanner (slice thic kness = 11.7 600 microns). The scans were segmented using VGStudioMax (Volume Gra phics GmbH, Germany). This process generates a stereolithography (STL) surface me sh of the three-dimensional geometry from stacked DICOM images acquired by the CT scanner. Each STL was then imported

PAGE 72

61 into Geomagic Studio 6 (Geomagic Inc., USA). STLs of teeth from extant species were refined by repairing artifacts from the scanni ng process. Geomagic was also used to rebuild sections of the fossil teeth, as all three specimens were partially damaged by taphonomic processes. Finally, the refined STLs were processed into FE models using Strand7 (G & D Comp uting, Australia). Finite element models were built using a solid mesh composed of four-noded tetrahedral elements (47563 to 1087617 elements). All models were designated as static, linearly elastic, and isotropic. Biological materials are largely viscoelastic, and tooth materials have been shown to be anisotropic (Waters, 1980), however we assumed the above conditions for this study for the sake of model simplicity. Enameloid and dentine (osteodentine or orthodentine) were designate d in each model based on the CT scans, as the distribution of these materials is visibl e in the DICOM images. The elastic modulus (E) for each material was taken from th e nanoindenation testing described above. Poissons ratio ( v ) is also required for FEA, but is unknown for shark enameloid or dentine. Instead, for both dentines and enam eloid, the Poissons ratio for mammal enamel and dentine were used ( v = 0.3 for both materials) (Waters, 1980). Lastly, boundary conditions were applied to prevent the m odel from moving through space when loaded, which is a mathematical requirement of FEA. For all tooth models, all external nodes on the tooth base were designated as fixed, which prevented displacement about these nodes. In reality, shark teeth are anc hored to the jaw with collagenous Sharpeys fibers, which allows some cusp displacement in the linguallabial plane, though the mechanics of shark tooth attachment and cusp rotation are not clear (Moss, 1970). As it has been hypothesized that this flexible attachment is a stress dissip ater (Frazzetta, 1994; Powlik,

PAGE 73

62 1995), modeling tooth attachment as fixe d provides an upper bound to possible mechanical scenarios. Five loading regimes were used (Figur e 2.3): (1) Distributed load on tooth tip to represent initial punc ture into a prey item (hereafter re ferred to as puncture); (2 & 3) Distributed load on each cutting edge to represent unidirectional draw, which would occur during head shaking behavior (draw) ; (4 & 5) Distributed load on labial & lingual cusp faces (respectivel y) to represent tooth impa led on struggling prey item (holding). For all loading regimes, a total di stributed load of 10 kN was used. Previous studies have shown bite forces for large elasmobranchs approaching this value ( Sphyrna mokarran, 4.3 m TL: 6080 N (Mara et al, Unpublished data); Carcharodon carcharias 6.4 m TL: 9320 N anterior, 18216 N posterior (Wro e et al., 2008)). Visual inspection of effective Von Mises stress dist ributions produced by FEA was used to identify sites of possible failure, which function as indicators of decreased structural strength. The use of Von Mises stresses is widely accepted for FE studies of biological structures (Dumont et al., 2005; McHenry et al., 2007; Rayfield, 2007; Moreno et al., 2008; Wroe et al., 2008). All procedures were in accordance with Univ ersity of South Florida Institutional Animal Care and Use Committee protocol number T3195. Results Nanoindentation Enameloid of S. tiburo has a mean hardness of 3.53 GPa and a mean Youngs modulus of 68.88 GPa, while enameloid of C. taurus had a hardness and Youngs modulus of 3.20 GPa and 72.61 GPa, respectively (Table 2.2). There was no significant difference

PAGE 74

63 between the two species for the material properties of enameloid (E: t=-1.680, p= 0.131; H: t=-2.051, p=0.074). Both hardness and Youngs modul us were higher for osteodentine ( C. taurus H=1.21, E=28.44) than for orthodentine ( S. tiburo, H=0.97, E=22.49 ) (E: t=4.763, p=0.001; H: t=-3.151, p=0.014) (Table 2.2). Finite Element Analysis When loaded in puncture, most teeth, regardless of morphology, showed the same general stress pattern. For al l teeth, the highest Von Mises stress magnitudes occurred during puncture. This stress was primarily concentrated at the site of loading, the cusp apex, and rapidly dissipated away from the apex (Fig. 2.4, Appendix B). Little stress is concentrated near the tooth base. Two teet h did not follow this general pattern. The Hexanchus griseus tooth had stress concentrated at th e notches between the cusplets, but still far from the base (Fig. 2.4d). The FE model for Sphyrna mokarran showed stress extending down the distal cutting edge to the notch, with a small amount of stress reaching the tooth base (Fig. 2.4c). During draw, many teeth tended to have st ress distributions that mirror those of a cantilever beam (Fig. 2.5, Appendix B). Stress tended to be concentr ated on the cutting edges, with the center largely uns tressed, akin to a neutral axis. This pattern is apparent in the models of Carcharhinus limbatus, Carcharhinus leucas, Carcharodon carcharias Negaprion brevirostris both Isurus oxyrinchus Cladodus sp., Hybodus sp., and Scymnodon ringens. The individual cusps of Xenacanthus compressus displayed stress patterns consistent with those of cantilever b eams. However, stress was also concentrated between the two main cusps, surrounding the median cusplet (Fig. 2.5c).

PAGE 75

64 In addition to stress concentrations on the mesial cutting edge, many of the tooth models that had notches on the distal cutti ng edge tended to have stress concentrated there (Fig. 2.5d, Appendix B). Stress was ofte n also concentrated in the space between the main cusp and cusplets or heels. Th is overall pattern was seen in teeth of C. leucas, Galeocerdo cuvier, H. griseus, Hybodus sp., Prioncae glauca, S. ringens, S. mokarran and the lateral tooth of I. oxyrinchus Many of these same teeth had stress concentrations at the dentine-enamel junction (DEJ) at the base of the crown, including C. leucas, C. limbatus, G. cuvier, P. glauca, and S. ringens (Fig. 2.5d, Appendix B) Overall, the holding models showed stress concentrated in a beam-like pattern; stress occurred on the lingual and labial cusp faces toward the base, with the neutral axis along the cutting edges (Fig. 2.6, Appendix B). Additional stress concentrations occurred along the DEJ for C. leucas, C. limbatus, Cladodont sp (Fig. 2.6a) G. cuvier, H. griseus, N. brevirostris, P. glauca (Fig 2.6b), S. mokarran, and the anterior tooth of I. oxyrinchus. Carcharodon carcharias, H. griseus, and S. ringens also had stress concentrated across the tooth base. Discussion While many studies of shark tooth functional morphology have focused on cutting mechanics, performance, and general morphology, the mechanical behavior of the tooth itself has been largely ignored. As teeth are subjected to loads during feeding, they undergo stress, strain, and poten tially failure. Both performa nce and structural strength are subject to natural selec tion. Therefore, to truly understand the evolution of shark teeth, both prey processing abil ity and structural parameters must be considered. Studies

PAGE 76

65 on elasmobranch teeth in particular have the potential to provide insight into the evolution of feeding in vertebrates in gene ral, as chondrichthyans are basal gnathostomes (Carroll, 1988). Nanoindentation While the values of hardness and Young s modulus are the first reported for dental materials of sharks and all fishes, resp ectively, there are some caveats. It has been shown that specimen storage, including chemi cal dehydration, influences the results of nanoindentation of mammalian teeth (Habelit z et al., 2002; Guidoni et al., 2006). While the samples in this study were simply air-dried, it is likely that this may have affected the results. Both enameloid and dentine are vi scoelastic, anisotropic materials, and the results may vary depending on the axis of lo ading (Rasmussen et al., 1976; Kinney et al., 2003; Shimizu and Macho, 2007). However for human molar enamel, nanoindentation tests parallel and perpendicula r to the hydroxyapatite crysta llites resulted in only 1.5-3% difference in both Youngs modulus and hardness, less than the standa rd deviations seen within the data set (Braly et al., 2007). Similarily, Lepidosiren teeth were microindented along the long axis of the t ooth and transverse to the same axis, hardness of the petrodentine only differed by approximately 0.2 GPa (Currey and Abeysekera, 2003). It is possible that testing direction is negligib le due to the nano-scale of these indentation tests (Braly et al., 2007). Likewise, testing different regions of the dentine or enameloid may also result in differences in material properties. This has been shown by nanoindentation studies of human dentine (Hosoya and Marshall, 2005), enamel (Cuy et al., 2002; Ge et al., 2005),

PAGE 77

66 and the DEJ (Marshall Jr. et al., 2001). Neos elachian shark enameloid is composed of three separate layers. The outermost layer, the SLE, is thin and not organized into bundles. The next layer, the PLE, is composed of parallel bundles of fibrous enameloid; these bundles run normal to th e outer tooth surface. The inner layer, the TFE, is composed on interwoven, less organized bundles of enameloid (Gillis and Donoghue, 2007). Coupled with previous studies on these la yers demonstrating that different layers resist compressive and tensile forces differently (Preuschoft et al., 1974), testing different regions of enameloid should produce different results. While the tests were performed away from the enamel edge or DEJ, we coul d not differentiate between the three layers when picking indentation sites. Overall, both the hardness and Youngs modulus for shark enameloid and both dentines are similar to values for other ve rtebrates (Table 2.2). The hardness of tooth materials of only two other fishes, Lepidosiren paradoxa and Protopterus aethiopicus, have been studied; there is no Youngs modulus data for any fishes. The hardness of both osteodentine and orthodentine, as determin ed by this study, are 125 181% higher than the dentine of both lungfishes and 10 128% higher than that of mammals. While they do not have enamel, lungfish have petrodentine, a mineralized dentine that is thought to function as enamel does (Currey and Abey sekera, 2003). Shark enameloid is 7 35% harder than petrodentine, but falls within the range of mammalian enamel. This may be related to the microstructure of these tissu es. Peterodentine is composed of masses of crystallites arranged in a criss-cross fashion, but no layeri ng occurs (Ishiyama and Teraki, 1990). Like enameloid, mammalian enamel occurs in layers, though the number of layers and arrangement of the enamel prisms among layers within varies among taxa

PAGE 78

67 (Yamakawa, 1959; Gustafson and Gustafson, 1967; Dumont, 1995; Maas and Dumont, 1999; Martin et al., 2003). Amphibian and rep tilian enamel varies from nonprismatic to layered prismatic, however material propert ies for these tetrapods have not been measured (Poole, 1967b; Sato et al., 1992; Sato et al., 2005). A co mprehensive study of enamel microstructure and material propert ies among vertebrate groups may elucidate the relationship between enamel layers and hardness. Both hardness and Youngs modulus were higher for osteodentine ( C. taurus) than for orthodentine ( S. tiburo ). Microanatomy may contribute to this difference, as the arrangement of the tubules differs in each material. Orthodentine has tightly packed, parallel tubules. These tubul es radiate from the central pulp cavity towards the outer surface of the tooth (Goto, 1991) Osteodentine, by contrast, is composed of numerous vascular canals surrounded by concentric layers of dentine, similar to osteons in spongy bone. The arrangement of the dental osteons varies from branching and meandering to highly organized with parallel osteons (Lund et al., 1992). Chemical composition may also contribute to mechanical differences between the two tissues. Osteodentine and orthodentine differ in the amounts of cal cium, phosphorus, magnesium, and sodium; osteodentine contains more calcium than or thodentine, whereas orthodentine contains more phosphorus, magnesim, and sodium (L und et al., 1992). Studies on mammalian enamel and dentine suggest th at hardness and Youngs modulus are positively correlated with calcium, as it is in ot her calcified tissues, and phosphate content, while sodium content shows the opposite trend (Lefevre et al., 1976; Brear et al., 1990; Currey, 1998b; Mahoney et al., 2000; Cuy et al., 2002).

PAGE 79

68 Finite Element Analysis Shark teeth appear to be structur ally strong during puncture. Under the biologically releveant lo ads in this study, stress concentra tion occurred at the cusp apex for the majority of the tooth models load ed in puncture (Fig. 2.4, Appendix B). In general, a well-designed tooth should utilize most of the tooth to resist applied loads, with unstressed material deemed inefficient. However, this loading regime is equivalent to the initial stages of puncture, where the tooth is either just comp ressing or barely ruptured the prey item. Stress is equivalent to force divided by area; as the tooth continues to puncture the prey item, more of the tooths surface ar ea will contact the prey item, lowering stress and utilizing more of the tooth to do so. Theref ore, these teeth appear to be well designed to resist stress during puncture. The puncture models of H. griseus and S. mokarran did not have stress concentrated only at the tooth apex (Fig. 2.4c,d). Stress was also concentrated in the spaces between the cusp and cusplets in the H. griseus model. Unlike many other notched teeth, the notch created by the main cusp and first distal cusplet is further from the tooth base on the H. griseus tooth and therefore closer to the load. It is possible that other notched teeth have enough material betw een the load and the notch to dissipate stress created by puncture, while the H. griseus tooth does not. Unde r extreme loads it is conceivable that material failure could occur at the region of the cu splets. Therefore, for puncture, the size and shap e of the apical half of the cusp may be more important, stresswise, than the shape of th e tooth in its entirety. The stress concentration for S. mokarran follows the distal cutting edge of the tooth. This is mechanically analogous to a column

PAGE 80

loaded in compression; the load is in line with the distal cutting edge and the stress distribution follows the expected load path. Many of the teeth loaded in draw and holding showed stress distributions consistent with failure in cantilever beams (Gere, 2004) (Fig. 2.5 & 2.6, Appendix B). Stress concentrations may be used as an indicator of possible material failure for these loading regimes. Tooth models showed stress concentrations occurred either at the DEJ or on the lower half of the cusp. Based on beam theory, failure in a cantilever beam should occur at the location of the maximum bending stress ( bend ), which is equal to the bending moment (M) divided by the section modulus (S) of the beam (Gere, 2004). The section modulus is given by Eq. (4): cIS (4) where I is the second moment of area and c is the extreme fiber length, the greatest distance from the neutral axis (Gere, 2004). A prismatic cantilever beam loaded at the free end should first fail at the base if mechanically sound, as S stays constant and M increases toward the base. Teeth are non-prismatic; that is, the cross-sectional shape is not constant. The location of the maximum bending stress will then also depend on a function related to the rate of cross-sectional shape change (Gere, 2004). The faster the section modulus increases, the further from the tooth base bend will be located. For example, the stress concentrations on the anterior I. oxyrinchus tooth in draw and holding are not located at the DEJ; instead they are located where the cusp becomes lingo-labially 69

PAGE 81

70 thicker. Consequently, failure of these teeth s hould occur not at their base but at the point where the cusp suddenly thickens; this is wh ere stress concentrations were located in these tooth models. Five teeth in draw did not have stress di stributions as predicted from beam theory, but did for holding: G. cuvier, H. griseus, Hybodus sp., P. glauca, and S. mokarran (Fig. 2.5 & 2.6, Appendix B). Geometrically, they do not qualify as beams in this loading regime, as their length (height of the cusp) is less than twice their width (mesodistal distance across the cusp). When the load was applied in holding, the width dimension changed to the lingo-labial th ickness of the cusp, and under this loading regime stress concentrations were consistent with a well-designed cantilever beam. Stress concentrations occur at the apex where th e loads were applied and at the notch for G. cuvier where there is relatively less material. While the notch is a potential site of weakness in the tooth, it is functionally impor tant. As sharks engage in head shaking behavior, as mimicked by loading the mode ls in draw, the notch may serve as stress concentrator on tough materials similar to a pa per cutter, helping to shear the material (Motta, 2004). Based on the analyses presented here, te eth cannot be placed into functional morphotypes based on stress distributions that occur during puncture, draw, and holding. In all loading cases, the extant and extinct teeth apparently are not limited by structure. No tooth is structurally stronger under one lo ading condition versus another, indicating that tooth failure may not occur often under these loading conditions. The only reported discussion of tooth breakage in sharks is th at narrow-based biconvex cutting teeth, such

PAGE 82

71 as those of I. oxyrinchus are commonly broken, but no da ta is given to support the statement (Williams, 2001). Despite this apparent over-engineering, sharks continuously replace their teeth throughout their lifetime. Tooth replacement rate s vary among taxa, varying from a week to every three months (Moss, 1967; Wass, 1973; Reif et al., 1978; Luer et al., 1990). Historically, this has not been the rule. In general, chondricthyans (including cladodont sharks), acanthodians, crossopterygians, and lu ngfishes retained their teeth (Reif, 1982; Carroll, 1988; Williams, 2001). It has been s uggested that the evolution of cutting teeth in sharks was not possible until a pattern of regular tooth replacement occurred, which is necessary to maintain effective teeth b ecause breakage is common in narrow-based cutting teeth (Williams, 2001). Among terrestrial mammalian predators, especially in the conical narrow canine teeth, wear and breakag e is common and problematic as teeth do not heal as bone does (Van Valkenbu rgh and Ruff, 1987; Van Valkenburgh, 1988). However, based on the results of this study, I propose that teeth are replaced frequently because of tooth wear, not tooth failure. Pred ators with sharp teeth tend to have lower bite forces than those with conical teeth, su ch as crocodilians and patherine cats (Wroe et al., 2008); comparatively, sharp teeth are subject ed to loads that may not be high enough to elicit failure. The results of performance testing in puncture (see Chapter 1) further support the tooth wear hypothe sis, though the effects of tooth wear on performance was not specifically tested. Teeth that contacted pr ey items with a relatively flat surface, such as P. glauca did not successfully puncture. Compre ssing the prey item during the initial stages of puncture creates compressive for ces beneath the tooth tip and tension on the surrounding prey tissues as the prey item bul ges around the tooth. These stresses together

PAGE 83

72 generate an effective shear stress which, if large enough, ruptures th e tissues (F razzetta, 1988). Wear on the tooth tip decreases sharpne ss, increasing the amount of surface area of the tooth-prey contact. This may decrease stress on the prey item to the extent that puncture (failure) is not possible. Similarly, wear may also affect the cutting edges. A razor blade will dull faster th an a pocket knife. Consequently, sharp blades tend to dull quickly. As the cutting edges are worn down, teeth may become less effective. Unfortunately, there are no data on rate of tooth wear or the effect of wear on performance. Conclusions The first goal was to determine the material properties for enameloid, osteodentine, and orthodentine via nanoindentation. While shark de ntines are harder than other vertebrate dentines, enameloid has similar hardness and Youngs modulus to mammalian enamel. This latter relationship may be due to sim ilar microstructures between shark enameloid and mammalian enamel. I then used finite element analysis to visualize stress distributions of fossil and extant shark teeth during puncture, unidirectional draw (cutting), and holding. The specific goals for th is portion of the study were to determine if tooth morphologies are more structurally strong during diffe rent loading regimes and to examine the role of morphological features, su ch as notches or cusp shape, on stress distribution. Under the loading and boundary co nditions, which are consistent with bite forces of large sharks, shark teeth are structurally strong. Teeth loaded in puncture have localized stress concentrations at the cusp apex that diminish rapidly away from the apex. When loaded in draw and holding, the majority of the teeth show stress concentrations

PAGE 84

73 consistent with mechanically sound canti lever beams. Notche s result in stress concentration during draw and may serve as a weak point; however they are functionally important for cutting prey during lateral he ad shaking behavior. As shark teeth are replaced regularly, it is proposed that the frequency of tooth replacement in sharks is driven by tooth wear, not tooth failure.

PAGE 85

Figure 2.1: Testing sites for nanoindentation on a cross-section of a generalized tooth cusp. The white area is dentine, grey is enamel, and black is the pulp cavity. X = approximate site of indentation. 74

PAGE 86

Figure 2.2: Phylogenetic tree of species included in this study, based on Compagno, 1988; Martin et al., 1992; Naylor, 1992; Shirai, 1996; Grogan and Lund, 2004. Branches are not drawn to scale. a = anterior, l = lateral. Line drawings show tooth used for each species. 75

PAGE 87

Figure 2.3: Loading regimes illustrated on bull shark tooth (Carcharhinus leucas, left) and lateral view of shortfin mako shark tooth (Isurus oxyrinchus, right). H = holding, L = lateral cutting (draw), P = puncture. 76

PAGE 88

77 Figure 2.4: Representative finite element models (FEMs) loaded in puncture. All views are of the labial side of the tooth. Arrow indicates detine-enamel junction. (a) Carcharhinus limbatus, (b) Carcharhinus leucas, (c) Sphyrna mokarran, (d) Hexanchus griseus.

PAGE 89

78 Figure 2.5: Representative FEMs loaded in draw on the distal cutting edge. All views are of the lingual side of the tooth. Arrow = dentine-enamel junction (DEJ). (a) Isurus oxyrinchus, anterior tooth, (b) Carcharodon carcharias, (c) Carcharhinus limbatus, (d) Galecoerdo cuvier, (e) Xenacanthus compressus.

PAGE 90

Figure 2.6: Representative FEMs loaded in holding on the lingual face of the tooth. The left view is the lingual side of the tooth; the right is the distal side of the tooth. (a) Cladodus sp., (b) Prionace glauca. 79

PAGE 91

Table 2.1: Tooth and jaw position information for each extant species in this study. Position = tooth family counting from the jaw symphysis. Specific jaw position for C. carcharias is not known. Species nJaw Position I. oxyrinchus (a) 3Lower 2 I. oxyrinchus (l) 3Upper 6 C. carcharias 3Upper "Anterior" S. mokarran 3Upper 6 G. cuvier 3Lower 3 C. leucas 3Upper 8 C. limbatus 3Upper 5 P. glauca 3Upper 7 N. brevirostris 3Lower 3 S. ringens 1Lower 1 H. griseus 1Lower 2 80

PAGE 92

Table 2.2: Material properties of tooth materials for vertebrates. H = hardness (megapascals, MPa), E = Youngs modulus (MPa). Values for shark enameloid, osteodentine, and orthodentine (bold) are means + standard error. All other values taken from (Waters, 1980; Currey, 1998a; Mahoney, 2000; Habelitz et al., 2001; Lutz, 2002; Currey and Abeysekera, 2003). SpeciesMaterialH (GPa)E (GPa)Lepidosiren paradoxaDentine0.43-Loxodonta africanaDentine0.437.7Monodon monocerosDentine0.538.9Protopterus aethiopicusDentine0.58-Bos taurusDentine0.617.8Capreolus capreolusDentine0.63-Rattus rattusDentine0.88-Homo sapiensDentine0.9219.89Sphyrna tiburoOrthodentine0.97 + 0.0322.49 + 0.08Carcharias taurusOsteodentine1.21 + 0.0728.44 + 0.99Lepidosiren paradoxaPetrodentine2.49-Bos taurusEnamel3.0073Protopterus aethiopicusPetrodentine3.12-Carcharias taurusEnameloid3.20 + 0.0972.61 + 2.11Capreolus capreolusEnamel3.23-Rattus rattusInner Enamel3.52-Sphyrna tiburoEnameloid3.53 + 0.5268.88 + 0.67Macaca mulattaEnamel3.63-Homo sapiensEnamel3.987.5 81

PAGE 93

82 Chapter 3: Evolutionary Relationships Between Shark Tooth Morphology and Ecology Abstract The existence of a relationship between morphology, ecology, and behavior is the central tenet of ecomorphology, which implies that th e evolution of morphology and ecology are tightly correlated. Shark t ooth morphotypes and their pres umed functions present an interesting test of this paradigm, as extant shark teeth encompass a variety of shapes. Often these morphotypes are used to predict ecology in the ab sence of other data, despite the fact that these rela tionships have not been tested. Ther efore the goal of this study is to employ phylogenetic comparative methods to te st, within an evolutionary context, whether a relationship exists between shark tooth morphology and ecology. Based on ecomorphological principles, I hypothesized that diet and habitat are predictive of shark tooth shape. For each of 44 extant shark spec ies, a series of morphometric measurements were taken on teeth on the right side of the upper and lower jaws of up to five individuals. These measurements were used to calculate quantit ative tooth morphology characters, including cusp aspect ratio, notch an gle, cusp inclination indices, and percent of tooth base overlap. Data about ecology and di et were taken from the literature. I then used phylogenetically-informed least square s regression and pairwise comparisons to determine whether habitat and diet were predictive of tooth shape. While some

PAGE 94

83 significant relationships were identified between aspects of tooth morphology and diet and habitat, few relationships and no clear pa ttern emerged. This suggests that the aspects of shark tooth morphology measured here ar e not related to ecology. The lack of a relationship between tooth morphology and ecolo gical variables has implications for the paleobiology of sharks, where assumptions ab out diet and evolution are made based on tooth shape due to the paucity of fossilized dietary information. Introduction That a relationship exists between morphology, ecology, and behavior is the central tenet of ecomorphology (Williams, 1972; Karr and James, 1975; Bock, 1994). This implies that evolution of morphology and ecology are tightly correlated (Losos, 1990), resulting in the predictive power of one to the ot her (Weins and Rotenberry, 1980; Motta and Kotrschal, 1992). Aspects of dental mo rphology has been correlated with ecology primarily for mammals (Hylander, 1975; Va n Valkenburgh and Ruff, 1987; Richard and Dewar, 1991; Janis, 1995; Freeman, 2000; Evans and Sanson, 2003; Evans et al., 2005; Cuozzo and Yamashita, 2007; Evans et al., 2007), though some studies exist for other vertebrates as well (Patchell and Shine, 1986; Turingan, 1994; Pete rson and Winemiller, 1997; Jackson et al., 1999; Herrel et al., 2004; Jackson and Fritts, 2004). For example in lacerterid lizards, omivores tend to have blunter, wider teeth with more cusps when compared to insectivores (H errel et al., 2004); snakes a nd legless lizards that are durophagous tend to have hinged teeth (Patch ell and Shine, 1986; Jackson et al., 1999; Jackson and Fritts, 2004). Loricarid fishes that sc rape substrates to feed tend to have large spatulate teeth, while those that injest fine detritus have rudimentar y teeth (Delariva and

PAGE 95

84 Agostinho, 2005). In the tetraodontiform fishes, tooth morphology is related to diet; for example, those with blunt, robust teeth te nd to be durophagous, wh ile those with thin, sharp teeth are planktivorous (Turingan, 1994). Shark tooth morphotypes and their presumed functions present an interesting test of this ecomorphological paradigm. Extant shark teeth encompass a wide variety of shapes, including teeth with tr iangular serrated cusps, obliq ue serrated and non-serrated cusps, notched serrated cusps, non-serrate d recurved cusps, multicusped teeth, and flattened tooth pavements. The trend has b een to divide teeth into morphotypes and assign qualitative predictive functions wit hout any biomechanical testing (Peyer, 1968; Cappetta, 1987; Motta, 2004). Small teeth wi th lateral cusplets are characterized as suited for clutching or seizing. These are f ound within the Orectolobiformes, Squatinidae, and Scliorhinidae. Tearing or puncturing teeth have narrow, tall cusps and are usually not serrated; examples include teeth of the lamniforms Isurus and Mitsukurina Teeth whose crowns are lingo-labially flatte ned and widen towards the base are designated as cutting teeth; many of the Carcharhinidae fall into th is category. Molariform teeth, such as those found in the horn sharks and many batoids, fa ll into crushing and grinding categories (Cappetta, 1987; Motta, 2004). Inevitably, links between these func tional morphotypes and ecology are made. For example, sharks with clutching teet h are often benthic or benthopelagic and presumably seize elusive midwater prey. El asmobranchs with crushing dentition prey upon shellfish, small fishes and cephalopods and tend to be bent hic or benthopelagic (Cappetta, 1987). Those with grinding denti tions are usually benthic and feed upon hard prey (Reif, 1976; Nobiling, 1977; Cappetta, 1987; Summers et al., 2004). This leads to

PAGE 96

85 predictions about ecology for i ndividual species where informa tion about diet and habitat are lacking. For example, Chlamydoselachus anguineus the frilled shark, is a poorly known deepwater shark. It has been predicte d that this shark f eeds on cephalopods and bottom associated fishes based on its tooth morphology (Compagno, 2001). Chaenogaleus macrostoma has a clutching dentition, and it is assumed that it preys on small fishes, cephalopods, and crustaceans (Compagno, 1984b). The same tendency to predict ecology and function from tooth morphology exists for individual fossil chondric hthyan teeth. The cartilaginous na ture of the skeleton leads to a fossil record composed almost entirely of teeth. Inferences about ecology are thus made from tooth morphology (Peyer, 1968; Cappetta, 1987; Williamson et al., 1993; Cicimurri, 2000, 2004; Stahl and Parris, 2004). For example, the Jurassic shark Sphendous has high crowned teeth (tearing type) and has been hypothesized to eat softbodied invertebrates, while cochliodont holocephalans are hypothesized to be durophagous based on their molariform dentition (Peyer, 1968; Cappetta, 1987). Inferences about shark evolution are also made based on tooth morphology and often cite aspects of ecology as a driving force of e volution (Schaeffer, 1967; Maisey, 1982). It has been hypothesized that selection for larger bladed teeth occurred with a change to a macrophageous diet (Williams, 2001); in ot her words, tooth morphology is determined by diet. While connections between shark toot h morphology, ecology, and evolution are often cited, the existence of a firm rela tionship between tooth morphology and ecology has not been rigorously tested. Based on eco morphological principles, I hypothesize that diet and habitat are predictive of shark toot h shape. While this relationship is often

PAGE 97

86 reversed when addressing ecology of individu al species, I am inte rested in a broader evolutionary picture. Therefore the goal of this study is to employ phylogenetic comparative methods to test, within an evolut ionary context, whether a relationship exists between shark tooth morphology and ecology. Materials and Methods Specimens The goal was to sample at least one species from each extant shark family, excluding filter feeders. In some cases, such as within the Carcharhinidae and Lamnidae, tooth morphology was quite varied among species; in which case up to four species per family were sampled. In total, 28 out of 34 families were sampled, totaling 44 species (Appendix C). A composite phylogeny was compiled using Mesquite 2.5 (Maddison and Maddison, 2008), based on primarily on Shirai (1996), w ith additional information for individual orders and families from other previously published phylogenies (Compagno, 1988; Martin et al., 1992; Nayl or, 1992; Shirai, 1996; Goto, 2001) (Fig. 3.1, 3.2). Branch lengths were assigned to unity as branch le ngths for the majority of these taxa are unknown. The composite tree was then expor ted to the PDDDIST program (Garland and Ives, 2000), which output a phylogenetic varian ce-covariance matrix for later analysis. Ecological information Major components for each species diet were gathered from the l iterature (Figure 3.2, Appendix D). Diet was then classified in tw o ways. The first was absence/presence of general prey categories: teleost, elasmobr anch, shrimp, crab, worms, cephalopod, hard

PAGE 98

87 mollusks & echinoderms, and mammal. The second classification grouped prey by general hardness: soft (worms, cephalopod), medium (teleost, elasmobranch, shrimp, mammal), and hard (hard mollusks & echinoderms, crab) (Kohlsdorf et al., 2008; Huber et al., in press) (Appendix E) The latter classification of pr ey is based on biomechanical studies on fish feeding and the different for ces required to punctu re prey of differing material properties (Hernandez and Motta, 1997; Korff and Wainwright, 2004; Chapter 1 of this dissertation). Both depth and habit were used to classify habitat for each shark species, following Musick et al. (2004) (Figure 3.2, Appendix E). For habit, species were designated as benthic, benthopelagic, or pelagic. Benthic shar k species are bottomassociated pump ventilators. Benthopelagic species split their time between the water column and the bottom and are typically ram ventilators. Pelagic species do not spend any time on the bottom and are also ram ventilators. For depth, species were designated as coastal, bathyal, or oceanic. Coastal spec ies live at depths between 0-200 m. Both bathyal and oceanic species are found in waters deeper than 200 m; however bathyal species are associated with th e ocean floor while oceanic sp ecies are not (Musick et al., 2004). Tooth morphometrics Tooth shape measurements were taken from twelve teeth from the functional row from one to five jaws for each species, specimens permitting (Appendix F). Only teeth from the right side of each jaw were used to account for any fluctuating asymmetry in tooth shape. For both the upper and lower jaws, c ounting from the jaw symphysis, teeth 1, 2, 4,

PAGE 99

88 5, 7, and 8 were measured (Fig. 3.3). This par ticular pattern allowed us to sample the anterior, middle, and posterior of each ja w. On each tooth, the following measurements based in part on Shimada (2005) and on characters previously cited to be related to tooth function (Cappetta, 1987; Frazz etta, 1988; Abler, 1992; Motta, 2004) were taken (Fig. 3.4): base-cusp width (BCW; maximum cusp wi dth), cusp height (CH; perpendicular from cusp apex to BCW), mesial cutting e dge length (MCL; distance between cusp apex and most mesial point of BCW), distal cut ting edge (DCL; distance between cusp apex and most distal point of BCW), notch widt h (NW; distance from notch across to MCL, parallel to BCW), and notch height (NH; pe rpendicular from cusp apex to NW). From these measurements, three ratios were calculate d and used in subsequent analyses: cusp aspect ratio (CAR; CH/BCW), apex aspect ratio (AAR; NH/NW), and cusp inclination (CI; MCL/DCL). The following measurements were also used in the following analyses: base overlap (BO; distance of either base overlap (+) or between bases (-) of adjacent tooth divided by the mean of BW for both ba ses), notch angle (NA; angle taken from cusp apex to notch to most distal point of BCW), and cusplet angle (CA; angle taken from cusp apex to notch to apex of lateral cusp lets or distal heel when present, CA = 0 if no cusplets present) Other aspects of tooth shape were quantified as disc rete states (Appendix E): number of lateral cusplets (LAT), presence/ab sence of molariform teeth (MT; 0 = absent, 1 = present), lingual-labial cusp curvature (LC; 0 = straight cusp, 1 = curved lingually, 2= recurved labially), and degree of serration (S E; 0 = none, 1 = weakly serrated (visible under a dissecting microscope), 2= strongly serra ted (visible to the na ked eye)) (Fig. 3.4).

PAGE 100

89 Statistical analysis To determine which ecological variables influe nce tooth shape, habitat (habit and depth) or diet (prey categories or ma terial categories) was set as the independent variables, and tooth shape parameters as the dependent variables. Because tooth shape variables were both continuous and discrete, we used two se parate analyses. The relationships between ecological and discrete tooth shape variables were analyzed using pairwise comparisons via Mesquite 2.5 (Maddison and Maddison, 2008). This technique investigates associations in character states between two binary characters via phylogenetically separate pairs, and avoids assumptions about ancestral states and branch lengths (Read and Nee, 1995; Maddison, 2000). Pairs were chosen such that taxa included in each pair differed in the states of their independent variable. To satisfy the binary requirement, discrete variables were transformed into sets of dummy variables before analysis. A significance level of 0.05 was used for all comparisons. Continuous tooth shape variables were analyzed using a series of multiple regressions via the Matlab progr am Regressionv2.m (Lavin et al., In press). Each tooth shape variable was regressed against habitat (h abit and depth) or di et using two separate regression models. The first was an ordinary least squares regression (OLS), which does not take phylogeny into account. The second regression was a phylogenetic least squares regression (PGLS), which used the phylogenetic variance-covariance ma trix to produce a weighted regression (Garland and Ives, 2000). To determine the best fit for each regression set, Akaike Information Criteri on (AIC) was used; the regression with the lower AIC is the better fit. If the PGLS had the lower AIC, then the data exhibits a phylogenetic signal (Lavin et al., In press) Absence of a phylogenetic signal suggests

PAGE 101

90 that the relationship is not due to phylogenetic inertia, but instead may be due to adaptation (Kohlsdorf et al., 2008). Because teet h might function as subsets or functional groups, the analysis was repeated for teeth 1 and 2 grouped as an ante rior subset, teeth 4 and 5 grouped as a lateral subset, and teeth 7 and 8 grouped as a poste rior subset, keeping lower and upper jaw teeth separate. Results Habit and depth Overall, there was no pattern of which tooth variables exhibited si gnificant relationships with habit and depth (Appendix G). For habit, only two individual t ooth variables showed a significant relationship. Th e notch angle (NA) for tooth 7 of the upper jaw was more acute in pelagic sharks than in benthic sharks. There was more base overlap (BO) between teeth 4 and 5 in the uppe r jaw for pelagic sharks than for benthic sharks, and this relationship exhibited a phyl ogenetic signal. Pairwise comparisons revealed no significant relationships. More individual relationships existed betw een tooth variables and depth than with habit (Appendix G). Notch angles were more acu te in coastal sharks than bathyal sharks for tooth 5 of the upper jaw, but the opposite was true of tooth seven of the upper jaw. Bathyal sharks had more tooth overlap for t eeth 7 and 8 in the lower jaw than coastal sharks, and this relationshi p exhibited a phylogenetic signa l. Aspect ratios also had relationships with depth. Coasta l sharks tended to have broade r teeth (lower cusp aspect ratio (CAR)) than bathyal sharks for tooth 1 of the lower jaw. The cusp apex tended to be narrower and taller (higher apex aspect ratio (AAR)) in teeth 2 and 5 of the upper jaw. Of

PAGE 102

91 these, only upper jaw tooth 2 exhibited a phylogenetic signal. Pa irwise comparisons showed no significant relationships between discrete tooth variables and depth. Because teeth might function as subsets or functional groups, the analysis was repeated for teeth 1 and 2 groupe d as an anterior subset, teet h 4 and 5 grouped as a lateral subset, and teeth 7 and 8 grouped as a posterior subset, keep ing lower and upper jaw teeth separate. Again, no overall pattern was f ound with regards to habitat (Appendix G). Bathyal sharks tended to have narrower, talle r cusps and apices (greater AAR and CAR) for the anterior (1,2) lower jaw teeth. For the same teeth, coastal-bathyal sharks also tended to have higher AAR. For posterior (7,8 ) lower jaw teeth, bathyal sharks also had teeth with greater CAR and more acute notch angles compared to coastal and oceanic species. Of these, the relationships between the anterior lower jaw group and AAR, and the posterior lower jaw group and CA R, exhibited a phyl ogenetic signal. Diet When major prey items were considered individually, no obvious pattern was found (Appendix H). Sharks that included cepha lopods as a major dietary component had smaller AARs (taller, narrower cusp apices) for tooth 5 of the upper jaw, while sharks that preyed on worms tended to have higher CARs (broader cusps) overall for tooth 7 of the upper jaw. Only the relationship between tooth 5 and cephalopods had a phylogenetic signal. BO between teeth 7 and 8 in th e upper jaw and 4 and 5 in both the upper and lower jaws was also related to diet and e xhibited a phylogenetic signa l. Teeth 4 and 5 of the lower jaw and 7 and 8 of the upper jaw te nded to have less overlap if the species

PAGE 103

92 included teleosts as a major dietary com ponent, while teeth 4 and 5 of the upper jaw overlapped less if shrimp was a major prey item. Cusp angle (CA) and notch angle ( NA) showed similarly complicated relationships. CA was smaller for teeth 1 a nd 2 of the upper jaw if elasmobranchs were included in the diet, while tooth 5 of the upper jaw had smaller CAs if worms were a major prey item. Notch angles for teeth 4, 7, and 8 of the upper jaw also showed a significant relationship with di et. Tooth 8 had smaller NAs if teleosts were included in the diet, while tooth 7 had larger NAs if el asmobranchs were a major dietary component. For tooth 4, NA was smaller if hard mollu sks and echinoderms were consumed, and higher if elasmobranchs or cephalopods were included in the diet. No phylogenetic signal was found for any of these relationships. Lastl y, cusp inclination (CI) was related to diet for tooth 1 of the lower jaw and teeth 5, 7, and 8 of the upper jaw. Tooth 1 had lower CIs (less inclined) if cephalopods were in the diet, and this was the only variable whose relationship with CI exhibite d a phylogenetic signal. Tooth 5 was more inclined (higher CI) if teleosts were in the diet, where tooth 7 was less inclined if shrimp was included in the diet. CI was lower for tooth 8 if shrimp was a major part of the diet, but higher if hard mollusks and echinoderms were prey items Pairwise comparisons identified no significant relationships. When prey items were pooled by general hardness, still no major pattern emerged (Appendix H). Tooth 1 of the upper jaw tended to have higher AAR (narrower, taller cusp apices) if hard prey was a part of the diet. Cusps were more inclined (higher CI) for tooth 5 of the upper jaw if sharks preyed on medium or hard prey. Less base overlap occurred between teeth 1 and 2 and teeth 4 a nd 5 of the upper jaw if medium prey items

PAGE 104

93 were a major dietary component. Of these rela tionships, only BO for teeth 4 and 5 of the upper jaw exhibited a phylogenetic signal. Pa irwise comparisons showed no significant relationships. When teeth were placed into functio nal groups, no general pattern emerged (Appendix H). Anterior (1,2) lower jaw teeth had lower CAR (s horter, broader cusps) if worms were a major dietary component, whereas the anterior upper jaw group had a higher CAR if elasmobranchs were a major prey item. Anterior upper jaw teeth were more inclined if mammals were included in the diet and had lower notch angles if elasmobranchs were included in the diet. La teral (4,5) upper jaw t eeth had more acute notches if teleosts were a major prey item, and posterior upper jaw teeth had less acute notches if teleosts and elasmobranchs were included in the diet. None of these relationships exhibited a phylogenetic signal. When prey was pooled by hardness, anterior lower jaw teeth were more inclined if hard prey was a major prey item; this relationship exhibited a phylogenetic signal. Discussion Ecological morphology assumes a relati onship between ecology/behavior and morphology and often assumes a predictive power of one to the other (Weins and Rotenberry, 1980; Losos, 1990; Motta and Kotrschal, 1992). The basis for this relationship is rooted in the idea that environmental constraints on ecology and morphology are parallel (Weins and Ro tenberry, 1980). Based on ecomorphological principles, morphology and ecology should be evolutionarily linked. Thus, I would predict that for shark teeth, ecology would be related to tooth morphology as it is for

PAGE 105

94 mammals (Van Valkenburgh, 1989; Sacco and Van Valkenburgh, 2004; Evans and Sanson, 2005; Lucas et al., 2008), reptiles (Her rel et al., 2004; Jacks on and Fritts, 2004; Kohlsdorf et al., 2008) or other fishes (Y amaoka et al., 1986; Blaber et al., 1994; Delariva and Agostinho, 2005). Often shark te eth are ascribed func tional roles without biomechanical testing, and th en ecology is predicted fr om tooth morphology. Some studies on elasmobranchs have shown a rela tionship between toot h shape and ecology, however these are largely on a species by species case (Nobiling, 1977; Tricas and McCosker, 1984; Cortes et al., 1996; Summers 2000; Estrada et al., 2006; Ramsay and Wilga, 2007; Dean et al., 2008). In this st udy, the correlated evolu tion of diet, habit, depth, and tooth morphology has been investigated using two comparative methods: phylogenetic pairwise comp arisons and phylogenetic l east square regression. While a number of significant isolated re lationships between tooth shape and diet, habit, and depth were found, there does not appear to be any overall pattern, phylogenetically or otherwise. Teeth from th e same region of the jaw did not have the same relationship with a particular variable. In many instances, a particular tooth would have a significant relationship with a hab itat or diet variable, and the adjacent tooth would not. For example, tooth 8 of the upper ja w was less inclined if shelled mollusks or echinoderms were included in the diet, but inclination of the adjacent tooth (tooth 7) did not have a significant relations hip with diet. The single ex ception included teeth 7 and 8 of the lower jaw, both of which were less in clined if shrimp was included in the diet. Some relationships contradicted each other. For example, notch angles were more acute in coastal sharks than bathyal sharks for tooth 5 of the upper jaw, but the opposite was true of tooth seven of the upper jaw. The large number of statistical tests may have led to

PAGE 106

95 spurious significant relationshi ps. All together, for tests i nvolving individual teeth, 198 pairwise comparisons and 396 multiple regres sions were performed, for a total of 594 tests. As the number of tests increases, the chances of making a Type I error, or rejecting H 0 when it is true, also increases (Quinn and Keough, 2002). Based on these findings, there is no suppor t for predicting the aspects of tooth shape in this study from diet or habita t in extant sharks. While this seems counterintuitive, lack of a relationship between morphology and ecology has been found numerous times (Weins and Rotenberry, 1980; Grossman, 1986; Block et al., 1991). Specifically, no relationship between toot h morphology and ecology has been found in other studies of teleosts (Motta, 1988; Li nde et al., 2004), reptiles (Reif, 1983) and mammals (Evans et al., 2007), indicating that th is is not an isolated case. It is possible that other components of the feeding apparatus are more re lated to ecology than these tooth measures. For example, while the ante rior teeth in sparid teleosts were not correlated with food type, the shape of the pr emaxilla was (Linde et al., 2004). However, this is contrary to many studies of tooth morphology and ecology in isolated elasmobranch species (Nobiling, 1977; Tricas and McCosker, 1984; Cortes et al., 1996; Summers, 2000; Estrada et al ., 2006; Ramsay and Wilga, 2007; Dean et al., 2008). It is likely that other factors may have either obs cured any existing relationship or contributed to a lack of a relationship between tooth morphology and ecology, including the choice of morphological parameters measured, classification of ecology, toot h biomechanics, and feeding behavior. While every effort was made to choose mo rphological characters that have been previously cited as contributing to tooth function (Nobiling, 1977; Cappetta, 1987;

PAGE 107

96 Frazzetta, 1988; Motta, 2004), ther e are other shape parameters that may be related to ecological factors such as diet. For example, the sharpness of the tooth tip was not measured, though studies on mammalian teeth have indicated that this directly impacts the ability for a given tooth to puncture prey items (Freeman and Weins, 1997; Evans and Sanson, 1998; Freeman and Lemen, 2007). The fl exible attachment of elasmobranch teeth to the jaw has also been linked functi onally to diet (Ramsay and Wilga, 2007; Dean et al., 2008). Similarly, hinged teeth have b een cited as an adaptation for durophagy in snakes (Stavitzky, 1981; Jacks on et al., 1999) The amount of tooth deflection could not be measured due to the nature of the specime ns used in this study, the majority of which were either dried jaws or fixed museum speci mens. Size may also be a factor leading to this lack of fit. Due to the disparity of an imal sizes, no direct meas ure of tooth size was included in this analysis; all linear measurem ents were used to calculate ratios (AAR, CAR, CI, BO). One of the shape parameters included in the desc ription of clutching dentition is small tooth size (Cappetta, 1987; Motta, 2004). The size of the predator and its feeding apparatus imposes constraints on diet and ecol ogy (Wainwright and Richard, 1995). The way in which ecology was classified may also contribute to the lack of ecomorphological relationship. The categorization of habitat into habit (benthic, benthopelagic, pelagic) and depth (costal, ocean ic, pelagic) may have been too coarse; for example, it has been shown that microhabita t is correlated with morphology in seagrass teleosts (Motta et al., 1995). Si milarly, our diet categorization may have lead to a lack of ecomorphological pattern. The dietary data us ed in this study was based on the most common prey items for each species and was categorized by taxonomy and presumed

PAGE 108

97 material properties. It has been argued th at prey should be organized by the functional challenges that they pose to the predator, which includes elusivity and prey size (Motta, 1988; Luczkovich et al., 1995; No rton, 1995; Norton et al., 19 95; Linde et al., 2004). The material properties of prey are just one of many possible functional challenges. Additionally, a predators choice of pr ey may not correspond with the upper limits of performance by the pr edator. The lacertilian lizard Varanus niloticus has blunt teeth, but does not specialize in hard prey; instead it eats a variety of organisms, including eggs, insects, and sma ll vertebrates; on the other hand, Ophisaurus apodus another lacertilian with blunt teeth, preys on snails (Reif, 1983). The teeth of carnivorous bears, such as polar bears Ursus maritimus are similar to omnivorous canids, such as African wild dogs Lycaon pictus ; this may be due to a simila rity in preference for prey smaller than their own body size which allows less reliance on specialized craniodental adaptation (Sacco and Van Valkenburgh, 2004). Heterodontus are known for their molariform teeth and inclusions of hard prey in the diet, however these sharks will feed on softer prey if it is available (Reif, 1983). When teeth were analyzed as functional groups (anterior, lateral, posterior), the same lack of a pattern was found and th e tests did not yiel d many significant relationships. Further confounding the relatio nship between ecology and performance, performance testing results indicated that not only did many shark teeth perform puncture and draw equally well, but also that teet h with different morphology often performed equally well (see Chapter 1). Thus, teeth w ith different shapes may be functionally equivalent, which would cont ribute to an absence of an ecomorphological relationship. The same tooth may also be used to process pr ey items with different material properties.

PAGE 109

98 For example, Chiloscyllium plagiosum which has typical small clutching teeth with multiple lateral cusplets will feed on cephalopods, teleosts, and crabs. Teeth remain erect when processing softer prey items, such as cephalopods and teleosts, and passively depress to crush the hard carapace of crabs (Ramsay and W ilga, 2007). While this is a dramatic example, many sharks include prey items of varyi ng material properties in their diet (Compagno, 1984a, b). Even a prey category such as teleosts will encompass prey items with various material properties. Sheepshead Archosargus probatocephalus, require more force for penetration by shark teeth than do ladyfish Elops saurus ; A. probatocephalus has thicker, larger scales than E. saurus (see Chapter 1). As shark teeth must fulfill their biological role in differe nt loading regimes (puncturing, cutting, etc.) with prey of different properties, it is un likely that a firm link between tooth shape and ecology exists. Behavior likely contributes to the lack of a relationship between tooth morphology and ecology. In addition to having a functional apparatu s to capture prey, predators also must have the ability to effectively use the apparatus via the appropriate prey capture behavior (Ferry -Graham et al., 2002). Morphology thus has been shown to be correlated not only with diet but with feeding behavior (Motta et al., 1995; Norton, 1995). For some butterflyfishes (Chaetodontidae) it has been suggested that how these fishes feed on coral (nipping versus scraping) ra ther than the fact th at they feed on coral tissues is more ecomorphologically important (Motta, 1988). In elasmobranchs, several behaviors reduce prey to manageab le pieces. The cookie cutter shark Isistius brasiliensis will remove plugs of tissue from marine mammals by biting and rotating the entire body (Jones, 1971; Compagno, 1984a; LeBeouf et al., 1987; Shirai and Nakaya, 1992),

PAGE 110

99 whereas the white shark Carcharodon carcharias uses head shaking (Frazzetta and Prange, 1987). Though both species possess broad, triangular teeth with visible serrations, there is a great difference in size and base overlap. The nurse shark Ginglymostoma cirratum which has small multi-cusped clutching teeth, will employ a spit-suck behavior to break up large prey (Matott et al., 1995), whereas carcharhinid sharks such as the bull shark Carcharhinus leucas which have broad serrated cutting teeth, will use head shaking (Frazzetta and Prange, 1987). When structures have multiple biological roles, they may not be optimized for any one role, making correlations between form and function difficult (Reif, 1983; Lauder, 1995; Koehl, 1996; Domenici and Blake, 2000). In addition to feeding, teeth in elasmobranchs are used for gripping females during mating, and this has also been shown to be related to tooth morphology in elas mobranchs (Springer, 1967; Compagno, 1970; Fedducia and Slaughter, 1974; McCourt a nd Kerstitch, 1980; Co mpagno, 1988; Kajiura and Tricas, 1996). Tooth morphology may theref ore not soley reflect adaptations to feeding. The lack of a relationship between these aspects of tooth morphology and ecological variables has imp lications for the paleobiology of sharks. The chondricthyan skeleton is composed primarily of cartilage, which does not fossilize well. This leads to a fossil record composed primarily of teeth. In formation about habitat can be obtained from rock facies and concurrent fossils, but few foss ils preserve direct evidence of shark diet (Zangerl and Richardson, 1963; Mapes and Ha nsen, 1984; Williams, 1990; Martill et al., 1994; Mapes et al., 1995; Schwimmer et al., 1997; Shimada and Hooks III, 2004; Kriwet et al., 2008). Instead, assumptions about diet are made based on tooth morphology, both

PAGE 111

100 with and without information from the surrounding rocks (Peyer, 1968; Zangerl, 1981; Lund, 1985, 1990; Whitenack et al., 2002; Ellio tt et al., 2004). While it may be tempting to maintain a link between ecology and tooth shape for Cenozoic fossil sharks that have teeth that are very similar to their living relatives, this analysis does not support this practice. This is especially true for tooth morphologies that are geometrically far removed from modern sharks, such as cladodont and xe nacanthid sharks. Perh aps a better strategy for interpreting feeding ecology would be to couple paleontological information on depositional environment and possible prey it ems with biomechanical testing such as performance testing and finite element analys is. This would give paleobiologists an idea of what teeth are actually capable of and thus refine hypotheses of function, biological role, and feeding ecology. Conclusions In this study, phylogenetic comparative me thods were employed to test whether diet and habitat are predictive of shar k tooth shape. Based on ecomorphological principles, I hypothesized that di et and habitat are predictive of shark tooth shape. While some significant relationships were identif ied between aspects of tooth morphology and diet and habitat, the relationshi ps were largely isolated and no clear pattern emerged. This suggests that the aspects of shark tooth mo rphology measured here are not related to ecology. Studies where ecological information is lacking have tended to use tooth shape to make assumptions about ecology, which this study does not support. Future studies incorporating prey captu re behavior and other tooth functio ns, such as use of teeth during mating, may elucidate the relationshi p between tooth morphology and ecology.

PAGE 112

101

PAGE 113

102 Figure 3.1: Phylogenetic tree of species included in this study, based on Compagno, 1988; Martin et al., 1992; Naylor, 1992; Shirai, 1996; Goto, 2001. Branches are not drawn to scale.

PAGE 114

Figure 3.2: Phylogenetic tree of species included in this study, with tooth morphology and ecology. B = benthic, BP = benthopelagic, P = pelagic, Ba = bathyal, C = coastal, O = oceanic, CB = coastal/bathyal, CO= costal/oceanic, E = elasmobranch prey, T = teleost, Sh = shrimp, Ce = cephalopod, M = mammal, Em = hard echinioderm/mollusk, w = worm. Prey items are color coded by hardness: Blue = soft, Red = medium, Green = hard. 103

PAGE 115

Figure 3.2 (continued). 104

PAGE 116

Figure 3.2 (continued). 105

PAGE 117

Figure 3.3: Upper jaw of Carcharhinus leucas illustrating tooth positions sampled and base overlap measurement (BO). S = symphysis. The same counting scheme applies to the lower jaw. 106

PAGE 118

Figure 3.4: Tooth measurements taken for evolutionary analysis. BCW = base-cusp width, BW = base width, CA = cusp angle, CH = cusp height, DCL = distal cutting edge, LC = lingo-labial curvature, MCL = mesial cutting edge, NA = notch angle, NH = notch height, NW= notch width. 107

PAGE 119

108 Conclusions The overall goal of this study was to i nvestigate the biomechanics and evolution of extant and extinct shark teeth. All part s of this study revolv e around tooth morphology, as tooth forms are often ascribed qualitativ e functions without any biomechanical testing and morphology is used to infer function and ecology in both extant and extinct sharks. For the first part of this study, I inves tigated the performance of three general categories of extant teeth, tear ing-type, cutting-type, and cu tting-clutching type, as well as three fossil morphologies, on a variety of prey items. The goals of this study were to: (1) Determine the forces necessary for individua l teeth to penetrate a variety of fish and crustacean prey representative of shark diets; (2) Determine what differences in penetration force and efficiency occur am ong tooth types; (3) Compare performance between different cutting regimes for a gi ven tooth morphology and (4) Determine which morphological aspects, if any, of tooth sh ape are predictive of tooth performance. Differences in puncturing performance occurr ed among different prey items, indicating that not all soft prey items are alike. Th e majority of teeth were able to puncture different prey items, and differences in puncture performance also occurred among tooth types; however, few patterns emerged. For ce to puncture was less than the maximum force that occurred during draw tests, how ever there were no differences between the maximum draw forces and maximum puncture forces. Few morphologi cal patterns were

PAGE 120

109 identified. In some cases, broader triangular teeth were less effective at puncturing than narrow-cusped teeth. Teeth from Galeocerdo cuvier, Prionace glauca,Hexanchus griseus, and Sphyrna mokarran were unable to puncture many of soft prey items. The flat surface of the tooth-prey contact may decrease stress on th e prey item to the extent that puncture (failure) is not possible. No morphologica l characteristics were correlated with maximum draw force. Many of the shark teeth in this study we re not only able to perform draw and puncture equally well, but tooth morphologies were functionally equivalent to each other. This does not s upport the use of tooth morphology to predict biological role. The second part of this study was a quanti tative study of structural mechanics and materials of shark teeth. The first goal was to determine the material properties for enameloid, osteodentine, and orthodentine vi a nanoindentation. While shark dentines are harder than other vertebra te dentines, enameloid has similar hardness and Youngs modulus to mammalian enamel. This latter relationship may be due to similar microstructures between shark enameloid a nd mammalian enamel. I then used finite element analysis to visualize stress distribu tions of fossil and extant shark teeth during puncture, unidirectional draw (c utting), and holding. The specific goals for this portion of the study were to determine if tooth morphol ogies are more structurally strong during different loading regimes and to examine the role of morphological features, such as notches or cusp shape, on stress distri bution. Under the loading and boundary conditions in this study, which are consistent with b ite forces of large sharks, shark teeth are structurally strong. Teeth loaded in puncture have localized st ress concentrations at the cusp apex that diminish rapidly away from the apex. When loaded in draw and holding,

PAGE 121

110 the majority of the teeth show stress concentr ations consistent with mechanically sound cantilever beams. Notches result in stress concentration during draw and may serve as a weak point; however they are functionally important for cutt ing prey during lateral head shaking behavior. As shark teeth are replaced regularly, it is proposed that the frequency of tooth replacement in sharks is driven by t ooth wear, not tooth failure. This is supported by the results from performance testing, as teet h that contacted prey with relatively flat surfaces (high surface areas) did not successful ly puncture; when teeth become worn, the surface area of the tooth-pr ey contact increases which may prohibit puncture. The third part of this study employed phylogenetic comparative methods to test whether diet and habitat are predictive of shark tooth sh ape. Based on ecomorphological principles, I hypothesized that di et and habitat are predictive of shark tooth shape. While some significant relationships were identif ied between aspects of tooth morphology and diet and habitat, the relationshi ps were largely isolated and no clear pattern emerged. This suggests that shark tooth mor phology, as measured here, is no t related to ecology. Studies where ecological information is lacking ha ve tended to use tooth shape to make assumptions about ecology, which this study does not support. Future studies incorporating prey captu re behavior and other tooth functio ns, such as use of teeth during mating, may elucidate the relationship between tooth morphology and ecology. Clearly, we have just begun to investig ate shark tooth biomechanics and the ecomorphological implications. Lacking clear ecomorphological and biomechanical correlations it is difficult to conjecture a bout the evolution of shark teeth and their biological roles without mo re information. This study primarily addressed the performance and biomechanics of isolated teet h, the majority of which are bladed. There

PAGE 122

111 are other morphologies of extant shark t eeth that were not in cluded in this study, including the molariform teeth of heter odontid sharks, the multicusped teeth of the prickly shark Echinorhinus cookei, and the unique tricuspid teeth of the frilled shark Chlamydoselachus anguineus for which there is no performance data. In reality, sharks teeth form a functional complex that work together. How teeth of the upper and lower jaws shear past each other and how teeth of the same jaw affect puncture and cutting during draw are unanswered questions. Includi ng these parameters may elucidate a better link between tooth morphology and performance. Furthermore, other aspects of teeth need to be addressed, including flexible toot h attachments (but see Ramsay and Wilga, 2007), tooth base overlap that may transmit forces to linked teeth (Frazzetta, 1988), lateral cusplets, and edge sharpness. While I have presented the first perfor mance and structural mechanics tests on fossil teeth, I have only sampled three teet h from over 400 million years of evolution. Even within broad generalizations of fossil shark tooth morphology (e.g. cladodont, xenacanth, hybodont), several different morphologi es exist that remain untested (Zangerl, 1981; Cappetta, 1987). Predicting ecology fr om tooth morphology is prevalent in paleontological studies, where th e cartilaginous nature of the skeleton leads to a fossil record composed primarily by isolated teeth. Th is analysis does not support this practice, especially for tooth morphologies that ar e geometrically far re moved from modern sharks, such as cladodont and xenacanthid sharks. Instead, a better strategy for interpreting feeding ecology would be to couple paleontological and geological information with biomechanics to generate hypotheses of paleoecology. This would give

PAGE 123

112 paleobiologists an idea of what teeth are actually capable of and thus refine hypotheses of function, biological role and feeding ecology.

PAGE 124

113 References Abler, W. L. 1992. The serrated teeth of tyra nnosaurid dinosaurs, and biting structures in other animals. Paleobiology 18:161-183. Anderson, P. S. L., and M. LaBarbera. 2008. Functional consequences of tooth design: effects of blade shape on energetics of cutting. Journal of Experimental Biology 211:3619-3626. Arnold, S. J. 1983. Morphology, performance and fitness. American Zoologist 23:347361. Barry, K. P. 2002. Feeding habits of blacktip sharks, Carcharhinus limbatus, and Atlantic sharpnose sharks, Rhizoprionodon terraenovae, in Louisiana costal waters. MS thesis/dissertation, Louisiana State University. Bemis, W. E., and G. V. Lauder. 1986. Mor phology and function of the feeding apparatus of the lungfish, Lepidosiren paradoxa (Dipnoi). Journal of Morphology 187:81108. Bennet, A. F. 1991. The evolution of activity capacity. Journal of Experimental Biology 160:1-23. Benton, M. J. 1997. Vertebrate Paleontol gy. Chapman and Hall, New York, 452 pp.

PAGE 125

114 Bethea, D. M., J. A. Buckel, and J. K. Carlson. 2004. Foraging ecology of the early life stages of four sympatric shark specie s. Marine Ecology Progress Series 268:245264. Biewiner, A. A. 1992. Overview of structural materials; pp. 1-20 i n A. A. Biewiner (ed.), Biomechanics Structures and Systems. A Practical Approach. I.R.L. Press, Oxford. Biknevicius, A. R., B. Van Valkenburgh, a nd J. Walker. 1996. Incisor size and shape: implications for feeding behaviors in sabe r-toothed "cats". Journal of Vertebrate Paleontology 16:510-521. Blaber, S. J. M., D. T. Brewer and J. P. Salini. 1994. Diet and dentition in tropical ariid catfishes from Australia. Environm ental Biology of Fishes 40:159-174. Block, W. M., L. A. Brennan, and R. J. Guitierrez. 1991. Ecomorphological relationships of a guild of ground-foraging birds in northern California, USA. Oecologia 87:449-458. Bock, W. J. 1980. The definition and recogni tion of biological adaptation. American Zoologist 20:217-227. Bock, W. J. 1994. Concepts and methods in ecomorphology. Journal of Bioscience 19:403-413. Bock, W. J., and G. von Walhert. 1965. Adaptation and the form-function complex. Evolution 19:269-299. Bonnet, E., and Y. Van de Peer. 2002. zt: a so ftware tool for simple and partial Mantel tests. Journal of Sta tistical Software 7:1-12.

PAGE 126

115 Bradford, E. W. 1967. Microanatomy and histochemistry of dentine; pp. 3-34 i n A. E. W. Miles (ed.), Structural and Chemical Organization of Teeth. Academic Press, New York. Brainard, E. L., and E. Azizi. 2005. Mu scle fiber angle, segment bulging, and architectural gear ratio in segmented musculature. Journal of Experimental Biology 208:3249-3261. Braly, A., L. A. Darnell, A. B. Mann, M. F. Telford, and T. P. Weihs. 2007. The effect of prism orientation on the indentation testi ng of human molar enamel. Archives of Oral Biology 52:856-860. Brear, K., J. D. Currey, C. M. Pond, and M. A. Ramsay. 1990. The mechanical preperties of the dentine and cement of the tusk of the narwhal Monodon monoceros compared with those of other minerali zed tissues. Archives of Oral Biology 35:615-621. Brett, C. E., and S. E. Walker. 2002. Pred ators and predation in Paleozoic marine environments. Paleontological Society Papers 8:93-118. Cappetta, H. 1986. Types dentaires adaptaif s chez les selaciens actuels et postpaleozoiques. Paleovertebrata 16:57-76. Cappetta, H. 1987. Chondrichthyes II. Handbook of Paleoichthyology. Gustav Fischer Verlag, New York, 191 pp. Carroll, R. L. 1988. Vertebrate Paleontology and Evolution. W. H. Freeman, New York, 698 pp. Castro, J. I. 1996. Biology of the blacktip shark, Carcharhinus limbatus, off the southeastern United States. Bulletin of Marine Science 59:508-522.

PAGE 127

116 Churcher, C. S. 1985. Dental functional morphology in the marsupial sabre-tooth Thylacosmilus atrox (Thylacosmilidae) compared to that of felid sabre-tooths. Australian Mammology 8:201-220. Cicimurri, D. 2000. Early Cretaceous elasm obranchs from the Newcastle Sandstone (Albian) of Crook County, Wyoming. The Mountain Geologist 37:101-107. Cicimurri, D. 2004. Late Cretaceous Chondricht hyans from the Carlie Shale (Middle Turonian to Early Coniacian) of th e Black Hills Region, South Dakota and Wyoming. The Mountain Geologist 41:1-16. Cigala-Fulgosi, F. 1995. Rare oceanic deep water squaloid sharks from the Lower Pliocene of the Northern Apennines (P arma Province) Italy. Bollettino della Societa Paleontologica Italiana 34:301-322. Compagno, L. J. V. 1970. Systematics of the genus Hemitriakis (Selachii: Carcharhinidae), and related genera. Pro ceedings of the California Academy of Sciences 38:63-98. Compagno, L. J. V. 1984a. FAO Species Catalo gue. Sharks of the World. An annotated and illustrated catalogue of shark species known to date, Part 1: Hexanchiformes to lamniformes. Food and Agriculture Organization of the United Nations, Rome. Compagno, L. J. V. 1984b. FAO Species Catalo gue. Sharks of the World. An annotated and illustrated catalogue of shark species known to date, Part 2: Carcharhiniformes. Food and Agricult ure Organization of the United Nations, Rome. Compagno, L. J. V. 1988. Sharks of the Order Carcharhiniformes. Princeton University Press, Princeton, 570 pp.

PAGE 128

117 Compagno, L. J. V. 2001. Sharks of the World: An annotated and illustrated catalogue of shark species known to date. Volume 2. Bullhead, mackerel and carpet sharks (Heterodontiformes, Lamniformes a nd Orectolobiformes). FAO Species Catalogue for Fishery Purposes. 2:1-269. Cortes, E. 1999. Standardized diet compositions and trophic levels of sharks. ICES Journal of Marine Science 56:707-717. Cortes, E., C. A. Manire, and R. E. Hueter 1996. Diet, feeding habi ts, and diel feeding chronology of the bonnethead shark Sphyrna tiburo in southwest Florida. Bulletin of Marine Science 58:353-367. Cundall, D., and A. Deufel. 1999. Striking pa tterns in booid snakes. Copeia 1999:868883. Cuozzo, F. P., and N. Yamashita. 2007. Impact of ecology on the teeth of extant lemurs: a review of dental adaptations, function, and life history; pp. 67-96 i n L. Gould and M. L. Sauther (eds.), Developments in Primatology: Progress and Prospects. Lemurs. Springer U.S., New York. Currey, J. D. 1998a. Mechanical properties of vertebrate hard tissues. Proceedings of the Institution of Mechanical Engineers, Part H 212:399-411. Currey, J. D. 1998b. The effect of porosity and mineral content on the Young's modulus of elasticity of compact bone. J ournal of Biomechanics 21:131-139. Currey, J. D., and R. M. Abeysekera. 2003. The microhardness and fracture surface of the petrodentine of Lepidosiren (Dipnoi), and of other mineralized tissues. Archives of Oral Biology 48:439-447.

PAGE 129

118 Cuy, J. L., A. B. Mann, K. J. Livi, M. F. Teaford, and T. P. Weihs. 2002. Nanoindentation mapping of the mechanical properties of human molar tooth enamel. Archives of Oral Biology 47:281-291. Dean, B. 1909. Studies on fossil fishes (sharks, chimaeroids, and arthrodires). Memoirs of the American Museum of Natural History 9:211-287. Dean, M. N., J. J. Bizzaro, and A. P. Summ ers. 2007. The evolution of cranial design, diet, and feeding mechanisms in batoid fishes. Integrative and Comparative Biology 47:70-81. Dean, M. N., J. B. Ramsay, and J. T. Sch aefer. 2008. Tooth reorie ntation affects tooth function during prey processing and toot h ontogeny in the le sser electric ray, Narcine brasiliensis Zoology 111:123-134. Delariva, R. L., and A. A. Agostinho. 2005. Relationship between morphology and diets of six neotropical loricarids. Journal of Fish Biology 58:832-847. Deufel, A., and D. Cundall. 1999. Do booids stab prey? Copeia 1999:1102-1107. Domenici, P., and R. W. Blake. 2000. Biomechanics in behaviour; pp. 1-17 i n P. Domenici and R. W. Blake (eds.), Bi omechanics in Animal Behaviour. BIOS Scientific Publishers Ltd., Oxford. Dudley, S. F. J., and G. Cliff. 1993. Sharks caught in the protective gill nets off Natal, South Africa. 7. The blacktip shark Carcharhinus limbatus (Valenciennes). South African Journal of Marine Science 13:237-254. Dullemeijer, P., and C. D. N. Barel. 1977. Functional morphology and evolution; pp. 83117 i n M. K. Hecht, P. C. Goody, and B. M. Hecht (eds.), Major Patterns in Vertebrate Evolution. Plenum Publishing Corporation, New York.

PAGE 130

119 Dumont, E. R. 1995. Mammalian enamel pris m patterns and enamel deposition rates. Scanning Microscopy 9:429-422. Dumont, E. R., J. Piccirillo, and I. R. Gro sse. 2005. Finite-element analysis of biting behavior and bone stress in the facial skeletons of bats. The Anatomical Record 283A:319-330. Dutta, H. M. 1992. Feeding mechanism in largemouth bass, Micropterous salmoides : cinematographic and electromyographic an alysis. Journal of Freshwater Biology 4:167-184. Elliott, D. K., R. B. Irmis, M. C. Hansen, and T. J. Olson. 2004. Chondrichthyans from the Pennsylvanian (Desmoinesian) Naco Fo rmation of central Arizona. Journal of Vertebrate Paleontology 24:268-280. Erickson, G. M., J. Catanese III, a nd T. M. Keaveny. 2002. Evolution of the biomechanical material properties of the femur. The Anatomical Record 268:115124. Estrada, J. A., A. N. Rice, L. J. Natanson, and G. B. Skomal. 2006. Use of isotopic analysis of vertebrae in reconstructi ng ontogenetic feeding ecology in white sharks. Ecology 87:829-834. Evans, A. R., and G. D. Sanson. 2005. Corresp ondence between tooth shape and dietary biomechanical properties in insectivor ous microchiropterans. Evolutionary Ecology Research 7:453-478. Evans, A. R., M. Fortelius, J. Jernvall, and J. T. Eronen. 2005. Dental ecomorphology of extant European carnivorans; pp. 223-232 i n E. Zadzinska (ed.), Current Trends in

PAGE 131

120 Dental Morphology Research: 13th In ternational Symposium on Dental Morphology. University of Lodz Press, Lodz. Evans, A. R., G. P. Wilson, M. Fortelius, a nd J. Jernvall. 2007. High-level similarity of dentitions in carni vorans and rodents. Nature 445:78-81. Evans, R. D., and G. D. Sanson. 1998. The e ffect of tooth shape on the breakdown of insects. Journal of Zoology, London 246:391-400. Evans, R. D., and G. D. Sanson. 2003. The t ooth of perfection: f unctional and spatial constraints on mammalian tooth shape. Bi ological Journal of the Linnean Society 78:173-191. Fedducia, A., and B. H. Slaughter. 1974. Sexual dimorphism in skates (Rajidae) and its possible role in differential nich e utilization. Evolution 28:164-168. Fenton, M. B., J. M. Waterman, J. D. Roth, E. Lopez, and S. E. Fienberg. 1998. Tooth breakage and diet: a compar ison of bats and carnivorans. Journal of Zoology, London 246:83-88. Ferry-Graham, L. A., D. I. Bolnick, and P. C. Wainwright. 2002. Using functional morphology to examine the ecology and evolution of speciali zation. Integrative and Comparative Biology 42:265-277. Fisher-Cripps, A. C. 2004. Nanoindentat ion. 2nd Edition. Springer, New York, 263 pp. Flint, M. H., A. S. Craig, H. C. Reilly, G. C. Gillard, and D. A. D. Parry. 1984. Collagen fibril diameters and glycosaminoglycan content of skins indices of tissue maturity and function. Connective Tissue Research 13:69-83.

PAGE 132

121 Frazzetta, T. H. 1966. Studies of the morphology and function of the skull in the Boidae (Serpentes). Part II. Morphology an d function of the jaw appartatus in Python sebae and Python molurus Journal of Morphology 118:217-296. Frazzetta, T. H. 1988. The mechanics of cutting and the form of shark teeth (Chondrichthyes: Elasmobran chii). Zoomorphology 108:93-107. Frazzetta, T. H. 1994. Feeding mechanisms in sharks and other elasmobranchs; pp. 31-57 i n V. L. Bels, M. Chardon, and P. Vanderwalle (eds.), Biomechanics of Feeding in Vertebrates: Advances in Comp arative and Environmental Physiology. Springer-Verlag, New York. Frazzetta, T. H., and C. D. Prange. 1987. M ovements of cephalic components during feeding in some requiem sharks (Carch arhiniformes: Carcharhinidae). Copeia 1987:979-993. Freeman, P. W. 1992. Canine teeth of bats (Mic rochiroptera): size, shape, and role in crack propagation. Biological Journal of the Linnean Society 45:97-115. Freeman, P. W. 2000. Macroevolution in Mi crochiroptera: Recoupling morphology and ecoogy with phylogeny. Evolutionary Ecology Research 2:317-335. Freeman, P. W., and W. N. Weins. 1997. Puncturi ng ability of bat caninie teeth: the tip; pp. 225-232 i n T. L. Yates, W. L. Gannon, a nd D. E. Wilson (eds.), Life Among the Muses: Papers in Honor of James. S. Findley. The Museum of Southwestern Biology, Albuquerque. Freeman, P. W., and C. A. Lemen. 2007. The trade-off between tooth strength and toothpenetration: predicting optimal shape of canine teeth. Journal of Zoology, London 273:273-280.

PAGE 133

122 Garland, T., Jr., and A. R. Ives. 2000. Using th e past to predict the present: Confidence intervals for regression equations in phylogenetic comparative methods. American Naturalist 155:346-364. Ge, J., F. Z. Cui, X. M. Wang, and H. L. Feng. 2005. Property variations in the prism and the organic sheath within enamel by nanoindentation. Biomaterials 26:3333-3339. Gembella, S., L. Ebmeyer, K. Hagen, T. Hannic h, K. Hoja, M. Rolf, K. Treiber, F. Vogel, and G. Weitbrecht. 2003. Evolutionary tr ansformations of my oseptal tendons in gnathostomes. Proceedings of the Royal Society of London, B. Gere, J. M. 2004. Mechanics of Material s. 6 Edition. Brooks/Cole-Thomas Learning, Toronto, 940 pp. Gilchrist, M. D., S. Keenan, M. Curtis, M. Cassidy, G. Byrne, and M. Destrade. 2008. Measuring knife stab penetration into skin simulant using a novel biaxial tension device. Forensic Science International 177:52-65. Gillis, J. A., and P. C. J. Donoghue. 2007. The homology and phylogeny of chondrichthyan tooth enameloid. Journal of Morphology 268:33-49. Goto, M. 1991. Evolutionary trends of t ooth structure in Chondricthyes; pp. 447-451 i n S. Suga and H. Nakahra (eds.), Mechanics and Phylogeny of Mineralization in Biological Systems. Springer-Verlag, Tokyo. Goto, T. 2001. Comparative anatomy, phylogeny a nd cladistic classification of the order Orectolobiformes (Chondricht hyes, Elasmobranchii). Memoirs of the Graduate School of Fisheries Science, Hokkaido University 48:1-100.

PAGE 134

123 Grogan, E. D., and R. Lund. 2004. The origin an d relationships of early Chondricthyes; pp. 3-32 i n J. C. Carrier, J. A. Musick, a nd M. R. Heithaus (eds.), Biology of Sharks and Thier Relatives. CRC Press, New York. Grossman, G. D. 1986. Food resource partitioning in a rocky intertidal fish assemblage. Journal of Zoology, London 1:317-355. Guidoni, G., J. Denkmayr, T. Schoberl, and I. Jager. 2006. Nanoindentation in teeth: influence of experimental conditions on local mechanical properties. Philosophical Magazine 86:5705-5714. Gustafson, G., and A.-G. Gustafson. 1967. Microa natomy and histochemistry of enamel; pp. 75-134 i n A. E. W. Miles (ed.), Structur al and Chemical Organization of Teeth. Academic Press, New York. Habelitz, S., S. J. Marshall, G. W. Ma rshall Jr., and M. Balooch. 2001. Mechanical properties of human dental enamel on the nanometre scale. Archives of Oral Biology 46:173-183. Habelitz, S., G. W. Marshall Jr., M. Bal ooch, and S. J. Marshall. 2002. Nanoindentation and storage of teeth. Jour nal of Biomechanics 35:995-998. Hebrank, M. R. 1980. Mechanical properties and locomotor functions of eel skin. BIological Bulletin 158:58-68. Hebrank, M. R., and J. H. Hebrank. 1986. The mechanics of fish skin: lack of an "external tendon" role in two te leosts. BIological Bulletin 171:236-247. Heithaus, M. R. 2001. The biology of tiger sharks ( Galeocerdo cuvier ) in Shark Bay, Western Australia: sex ratio, size distribution, diet, and se asonal changes in catch rates. Environmental Biology of Fishes 61:25-36.

PAGE 135

124 Hernandez, L. P., and P. J. Motta. 19 97. Trophic consequences of differential performance: ontogeny of oral jaw-crushing performance in the sheepshead, Archosargus probatocephalus (Teleostei, Sparidae). Journal of Zoology, London 243:737-756. Herrel, A., B. Van Hooydonck, and R. Van Damme. 2004. Omnivory in lacertid lizards: adaptive evolution or constraint? J ournal of Evolutionary Biology 17:974-984. Hoffmayer, E. R., and G. R. Parsons. 2003. Food habits of three shark species from the Mississsippi Sound in the northern Gulf of Mexico. Southeast Naturalist 2:271280. Hosoya, Y., and G. W. Marshall. 2005. The nano-hardness and elastic modulus of sound deciduous canine dentin and young premolar dentin preliminary study. Journal of Materials Science: Materials in Medicine 16:1-8. Huber, D. R., and P. J. Motta. 2004. Compar ative analysis of methods for determining bite force in the spiny dogfish Squalus acanthias Journal of Experimental Zoology 301A:26-37. Huber, D. R., C. L. Weggelaar, and P. J. Mo tta. 2006. Scaling of bite force in the blacktip shark Carcharhinus limbatus Zoology 109:109-119. Huber, D. R., M. N. Dean, and A. P. Su mmers. 2008. Hard prey, soft jaws, and the ontegeny of feeding mechanics in the spotted ratfish Hydrolagus colliei Proceedings of the Royal Society Interface 5:941-952. Huber, D. R., T. G. Eason, R. E. Hueter, and P. J. Motta. 2005. Analysis of the bite force and mechanical design of the feeding mechanism of the durophagous horn shark Heterodontus francisci Journal of Experime ntal Biology 208:3553-3571.

PAGE 136

125 Huber, D. R., J. M. Claes, J. Mallefet, and A. Herrel. in press. Is extreme bite performance associate with extreme mor phologies in sharks? Physiological and Biochemical Zoology. Hylander, W. L. 1975. Incisor si ze and diet in anthropoids with special reference to Cercopithecidae. Science 189:1095-1098. Ishiyama, M., and Y. Teraki. 1990. The fine structure and formation of hypermineralized peterodentine in the tooth plate of extant lungfish ( Lepidosiren paradoxa and Protopterus sp. ). Archives of Histology and Cytology 53:307-321. Iturralde-Vinent, M., G. Hubbell, and R. Rojas. 1996. Catalogue of Cuban fossil Elasmobranchii (Paleocene to Pliocene) a nd paleogeographic implications of their Lower to Middle Miocene occurrence. J ournal of the Geological Society of Jamaica 31:7-21. Jackson, K., and T. H. Fritts. 2004. Dentit ional specializations for durophagy in the Common Wolf snake, Lycodon aulicus capucinus Amphibia-Reptilia 25:247254. Jackson, K., G. Underwood, E. N. Arnold, and A. H. Savitzky. 1999. Hinged teeth in the enigmatic colubrid, Iguanognathus werneri Copeia 1999:815-818. Janis, C. M. 1995. Correlations between cran iodental morphology and feeding behavior in ungulates: reciprocal illumination be tween living and fossil taxa; pp. 76-98 i n J. J. Thomason (ed.), Functional Mo rphology in Vertebrate Paleontology. Cambridge University Press, Cambridge. Johansen, E. 1967. Ultrastruc ture of dentine; pp. 35-75 i n A. E. W. Miles (ed.), Structural and Chemical Organization of Teet h. Academic Press, New York.

PAGE 137

126 Jones, E. C. 1971. Isitius brasiliensis a squaloid shark, the probable of crater wounds on fishes and cetaceans. U.S. Fisheries Bulletin 69:791-798. Jones, M., E.H. 2008. Skull shape and feeding strategy in Sphenodon and other Rhynchocephalia (Diapsida: Lepidosauri a). Journal of Morphology 269:945-966. Kajiura, S. M., and T. C. Tricas. 1996. Seasona l dynamics of dental se xual dimorphism in the Atlantic Stingray, Dasyatis sabina Journal of Experimental Biology 199:2297-2306. Karr, J. R., and F. C. James. 1975. Eco-morphological configurat ions and convergent evolution of species a nd communities; pp. 258-291 i n M. L. Cody and J. Diamond (eds.), Ecology and Evolution of Communities. Belknap, Cambridge, Massachusetts. Kinney, J. H., S. J. Marshall, and G. W. Ma rshall Jr. 2003. The mechanical properties of human dentin: a critical review and re-evalu ation of the dental literature. Critical Reviews of Oral Biology and Medicine 14:13-29. Koehl, M. A. R. 1996. When does morphol ogy matter? Annual Review of Ecological Systems 27:501-542. Kohlsdorf, T., M. B. Grizante, C. A. Navas, and A. Herrel. 2008. Head shape evolution in Tropidurinae lizards: does locomotion cons train diet? Journal of Evolutionary Biology 21:781-790. Konow, N., and C. P. J. Sanford. 2008. Is a c onvergently derived mu scle-activity pattern driving novel raking behaviours in tele ost fishes? Journal of Experimental Biology 211:989-999.

PAGE 138

127 Korff, W. L., and P. C. Wa inwright. 2004. Motor pattern co ntrol for increasing crushing force in the striped burrfish ( Chilomycterus schoepfi ). Zoology 107:335-346. Kotrshal, K., and A. Goldschmidt. 1992. Morpho logical evidence for th e biological role of caniniform teeth in combtooth blennies (Blenniidae, Teleostei). Journal of Fish Biology 41:983-991. Kriwet, J., F. Witzmann, S. Klug, and U. H. J. Heidtke. 2008. First direct evidence of a vertebrate three-level trophic chain in the fossil record. Proceedings of the Royal Society of London, B 275:181-186. Lauder, G. V. 1980. The suction feeding mechanism in sunfishes ( Lepomis ): an experimental analysis. Journal of Experimental Biology 88:49-72. Lauder, G. V. 1991. Biomechanics and evolutio n: integrating physical and historical biology in the study of complex systems; pp. 1-20 i n J. M. V. Rayner and R. J. Wooten (eds.), Biomechanics in Evol ution. Cambridge University Press, Cambridge. Lauder, G. V. 1995. On the inference of function from structure; pp. 1-18 i n J. J. Thomason (ed.), Functional Morpholgy in Vertebrate Paleontology. Cambridge University Press, Cambridge. Lavin, S. R., W. H. Karasov, A. R. Ives, K. M. Middleton, and T. Garland, Jr. In press. Morphometrics of the avian small intestine, compared with non-flying mammals: a phylogenetic approach. Physiological and Biochemical Zoology. LeBeouf, B. J., J. E. McCosker, and J. He witt. 1987. Crater wounds on northern elephant seals: the cookiecutter shark strikes again. U.S. Fisheries Bulletin 85:387-392.

PAGE 139

128 Lefevre, R., R. M. Frank, and J. C. Vo egel. 1976. The study of human dentine with secondary ion microscopy and electron diffraction. Calcified Tissue Research 19:251-261. Linde, M., M. Palmer, and G.-Z. J. 2004. Differe ntial correlates of diet and phylogeny on the shape of the premaxilla and anterior tooth in sparid fishes (Perciformes: Sparidae). Journal of Evolutionary Biology 17:941-952. Long, D. J. 1993. Late Miocene and Early Pl iocene fish assemblages from the northcentral coast of Chile. Tertiary Research 14:117-126. Losos, J. B. 1990. Ecomorphology, performance capability, and scaling of West Indian Anolis lizards: an eovlutionary anal ysis. Ecological Monographs 60:369-388. Lucas, P. W. 1982. Basic princi ples of tooth design; pp. i n B. Kurten (ed.), Teeth: Form, Function, and Evolution. Columbia University Press, New York. Lucas, P. W., C. R. Peters, and S. R. A rrendale. 1994. Seed-breaking forces exerted by orang-utans with their teeth in captiv ity with a new technique for estimating forces produced in the wild. American Journal of Physical Anthropology 94:365378. Lucas, P. W., P. J. Constantino, and B. A. Wood. 2008. Inferences re garding the diet of extinct hominins: structural and functi onal trends in dental and mandibular morphology within the hominin clade. Journal of Anatomy 212:486-500. Lucifora, L. O., R. C. Menni, and A. H. Escalante. 2001. Analysis of dental insertion angles in the sand tiger shark, Carcharias taurus (Chondrichthyes: Lamniformes). Cybium 25:23-31.

PAGE 140

129 Luczkovich, J. J., S. F. Norton, and R. G. Gilmore. 1995. The influence of oral anatomy on prey selection during the ont ogeny of two percoid fishes, Lagodon rhomboides and Centropomus undecimalis Environmental Biology of Fishes 44:79-95. Luer, C. A., P. C. Blum, and P. W. Gilbert. 1990. Rate of tooth replacement in the nurse shark, Ginglymostoma cirratum Copeia 1990:182-191. Lund, R. 1985. Ecomorphology of the Chondrict hyes from the Bear Gulch Limestone (Lower Carboniferous) of Montana; pp. 481-491 i n J. T. Dutro, Jr. and W. Hermann (eds.), 9th International Car boniferous Congress. International de Stratigraphie et de Geologie du Carbonifere, Washington, D.C. Lund, R. 1990. Chondrichthyan life history styles as revealed by the 320 million years old Mississippian of Montana. Envi ronmental Biology of Fishes 27:1-19. Lund, R., P. Bartholomew, and A. Kemp. 1992. The composition of the dental hard tissues of fishes; pp. 35-71 i n P. Smith and E. Tchernov (eds.), Structure, Function, and Evolution of Teeth. Freund Publishing House Ltd., London. Lutz, W. 2002. Differential tooth wear in a roe deer buck ( Capreolus capreolus Linne 1758). Zeitschrift fur Ja gdwissenschaft 48:194-202. Maas, M. C., and E. R. Dumont. 1999. Built to last: the structure, function, and evolution of primate dental enamel. E voultionary Anthropology 8:133-152. Maddison, W. P. 2000. Testing character corr elation using pairwi se comparisons on a phylogeny. Journal of Theoretical Biology 202:195-204. Maddison, W. P., and D. R. Maddison. 2008. Mesquite: a modular system for evolutionary analysis. 2.5. http://mesquiteproject.org.

PAGE 141

130 Mahoney, E. 2000. The hardness and modulus of el asticity of primary molar teeth: an ultra-microindentation study. J ournal of Dentistry 28:589-594. Mahoney, E., A. Holt, M. Swain, and N. Kilpatrick. 2000. The hardness and modulus of elasticity of primary molar teeth: an ultra-micro-i ndentation study. Journal of Dentistry 28:589-594. Maisey, J. G. 1982. The anatomy and relationships of Mesozoic hybodont sharks. American Museum Novitates 2724:1-48. Mapes, R. H., and M. C. Hansen. 1984. Penns ylvanian shark cephalopod predation: a case study. Lethaia 17:175-183. Mapes, R. H., M. S. Sims, and D. R. Boardmann II. 1995. Predation on the Pennsylvanian ammonoid Gonioloboceras and its implcations for allochthonous vs. autochthonous accumulations of goniat ites and other ammonoids. Journal of Paleontology 69:441-446. Marshall Jr., G. W., M. Balooch, R. R. Galla gher, S. A. Gansky, and S. J. Marshall. 2001. Mechanical poperties of the dentinoenamel junction: AFM studies of nanohardness, elastic modulus, and fractur e. Journal of Biomedical Materials Research Part A 54:87-95. Martill, D. M., M. A. Taylor, K. L. Duff, J. B. Riding, and P. R. Bown. 1994. The trophic study of the biota of the Peterboroug h Member, Oxford Clay Formation (Jurassic), UK. Journal of the Geological Society, London 151:173-194. Martin, A. P., G. J. P. Naylor, and S. R. Palumbi. 1992. Rates of mitochondrial DNA evolution in shark are slow comp ared to mammals. Nature 357:153-155.

PAGE 142

131 Martin, L. B., A. J. Olejniczak, and M. C. Maas. 2003. Enamel thickness and microstructure in pitheciin primates, with comments on dietary adaptations of the middle Miocene hominoid Kenyapithecus Journal of Human Evolution 45:351367. Matott, M. P., P. J. Motta, and R. E. Huet er. 1995. Modulation in feeding mechanics and motor pattern of the nurse shark Ginglymostoma cirratum Environmental Biology of Fishes 74:163-174. McCourt, R. M., and A. N. Kerstitch. 1980. Mating behavior and sexual dimorphism in dentition in the stingray Urolophus concentricus from the Gulf of California. Copeia 1980:900-901. McHenry, C. R., S. Wroe, P. D. Clausen, K. Moreno, and E. Cunningham. 2007. Supermodeled sabercat, predatory behavior in Smilodon fatalis revealed by highresolution 3D computer simulation. Pro ceedings of the National Academy of Sciences 104:10610-10615. Mertinene, R. A. 1982. The histology of the teeth of elasmobranchs. Paleontological Journal 16:74-82. Moreno, K., S. Wroe, P. D. Clausen, C. R. Mc Henry, D. C. D'Amore, E. J. Rayfield, and E. Cunningham. 2008. Cranial performance in the Komodo dragon ( Varanus komodoensis ) as revealed by high-resolution 3-D fintie element analysis. Journal of Anatomy 212:736-746. Moss, M. L. 1970. Enamel and bone in shark te eth: with a note on fibrous enamel in fishes. Acta Anatomy 77:161-187.

PAGE 143

132 Moss, S. A. 1967. Tooth replacement in the lemon shark, Negaprion brevirostris ; pp. 319-329 i n P. W. Gilbert, R. F. Mathewson, and D. P. Rall (eds.), Sharks, Skates, and Rays. John Hopkins University Press, Baltimore. Moss, S. A. 1977. Feeding mechanisms in sharks. American Zoologist 17:355-364. Motta, P. J. 1977. Anatomy and functional morphology of dermal collagen fibers in sharks. Copeia 1977:454-464. Motta, P. J. 1988. Functional morphology of th e feeding apparatus of ten species of Pacific butterflyfishes (Perciform es, Chaetodontidae): an ecomorphological approach. Environmental Biology of Fishes 22:39-67. Motta, P. J. 2004. Prey capture behavior a nd feeding mechanics of elasmobranchs; pp. 165-202 i n J. C. Carrier, J. A. Musick, a nd M. R. Heithaus (eds.), Biology of Sharks and Their Relatives. CRC Press, New York. Motta, P. J., and K. M. Kotrschal. 1992. Correlative, experimental, and comparative evolutionary approaches in ecomor phology. Netherlands Journal of Zoology 42:400-415. Motta, P. J., K. B. Clifton, L. P. Hern andez, and B. T. Eggold. 1995. Ecomorphological correlates in ten species of subtropical seagrass fish es: diet and microhabitat utilization. Environmental Biology of Fishes 44:37-60. Motta, P. J., R. E. Hueter, T. C. Tricas, A. P. Summers, D. R. Huber, D. Lowry, K. R. Mara, M. P. Matott, L. B. Whitenack, and A. P. Wintzer. 2008. Functional morphology of the feeding apparatus, feeding constraints, and suction performance in the nurse shark Ginglymostoma cirratum Journal of Morphology 269:1041-1055.

PAGE 144

133 Musick, J. A., M. M. Harbin, and L. J. V. Compagno. 2004. Historical Zoogeography of the Selachii; pp. 33-78 i n J. C. Carrier, J. A. Music k, and M. R. Heithaus (eds.), Biology of Sharks and Their Re latives. CRC Press, New York. Naylor, G. J. P. 1992. The phylogenetic rela tionships among requiem and hammerhead sharks: inferring phylogeny when thousands of equally most parsimonious trees result. Cladistics 8:295-318. Nobiling, G. 1977. Die Biomechanik des Kiefferapparates beim Stierkopfhai ( Heterodontus portusjacksoni = Heterodontus philippi ). Advances in Anatomy, Embryology, and Cell Biology 52:3-52. Norton, S. F. 1995. A functional approach to th e ecomorphological patterns of feeding in cottid fishes. Environmental Biology of Fishes 44:61-78. Norton, S. F., J. J. Luczkovich, and P. J. Motta. 1995. The role of ecomorphological studies in the comparative biology of fi shes. Environmental Biology of Fishes 44:287-304. Patchell, F. C., and R. Shine. 1986. Hinge d teeth for hard-bodi ed prey: a case of convergent evolution between snakes a nd legless lizards. Journal of Zoology, London (A) 208:269-275. Peterson, C. C., and K. O. Winemiller. 1997. On togenetic diet shifts and scale-eating in Roeboides dayi a Neotropical characid. Environmental Biology of Fishes 49:111118. Peyer, B. 1968. Comparative Odontology. Univer sity of Chicago Press, Chicago, 347 pp. Plotnick, R. E., and T. K. Baumiller. 2000. Inve ntion by evolution: functional analysis in paleobiology. Paleobiology 26:305-323.

PAGE 145

134 Poole, D. F. G. 1967a. Enameloid and enamel in recent vertebrates; pp. 111-150 i n A. E. W. Miles (ed.), Structural and Chemical Organization of Teeth. Academic Press, New York. Poole, D. F. G. 1967b. Phylogeny of tooth tissues: enameloid and enamel in Recent vertebrates, with a note on th e history of cementum; pp. 111-149 i n A. E. W. Miles (ed.), Structural and Chemical Organization of Teeth. Academic Press, New York. Porter, M. E., T. J. Koob, and A. P. Summers. 2007. The contribution of mineral to the material properties of vertebral cartilage from the smooth-hound shark Mustelus californicus Journal of Experime ntal Biology 210:3319-3327. Porter, M. E., J. L. Beltran, T. J. Koob, and A. P. Summers. 2006. Material properties and biochemical composition of mineralized vert ebral cartilage in seven elasmobranch species (Chondrichthyes). Journal of Experimental Biology 209:2920-2928. Powlik, J. J. 1995. On the geometry and mech anics of tooth position in the white shark Carcharodon carcharias Journal of Morphology 226:277-288. Pratt, H. L., and J. C. Carrier. 2001. A revi ew of elasmobranch reproductive behavior with a case study on the nurse shark, Ginglymostoma cirratum Environmental Biology of Fishes 60:157-188. Preuschoft, H., W.-E. Reif, and W. H. Mu ller. 1974. Funktionsanpassungen in Form und Struktur an Haifischzhnen. Z Anat Entwicklungsgesch 143:315-344. Purdy, R., V. P. Schnieder, S. P. Applegate, J. H. McLellan, R. L. Meyer, and B. H. Slaughter. 2001. The Neogene sharks, ra ys, and bony fishes from Lee Creek

PAGE 146

135 Mine, Aurora, North Carolina. Geology and paleontology of the Lee Creek Mine, North Carolina, III. Smithsonian Cont ributions to Paleobiology 90:71-202. Purdy, R. W. 1996. Fish teeth from the Pleistocene of Jamaica. Journal of Vertebrate Paleontology 16:165-167. Quinn, G. P., and M. J. Keough. 2002. Experime ntal Design and Data Analysis for Biologists. Cambridge Univer sity Press, New York, 537 pp. Ramsay, J. B., and C. D. Wilga. 2007. Morphology and mechanics of the teeth and jaws of white-spotted bamboo sharks ( Chiloscyllium plagiosum ). Journal of Morphology 268:664-682. Rana, R. S., K. Kumar, R. S. Loyal, A. Sahni K. D. Rose, J. Mussell, H. Singh, and S. K. Kulshrestha. 2006. Selachians from the Ea rly Eocene Kapurdi Formation (Fuller's Earth), Barmer District, Ra jasthan. Journal of the Ge ological SOciety of India 67:509-522. Rasmussen, S. T., R. E. Patchin, D. B. Sco tt, and A. H. Heuer. 1976. Fracture properties of human enamel and dentin. Jour nal of Dental Research 55:154-164. Rayfield, E. J. 2007. Finite element analys is and understanding the biomechanics and evolution of living and fossil organisms. Annual Review of Earth and Planetary Sciences 35:541-576. Read, A. F., and S. Nee. 1995. Inference fr om binary comparative data. Journal of Theoretical Biology 173:99-108. Reif, W.-E. 1976. Morphogenesis, pattern form ation and function of the dentition of Heterodontus (Selachii). Zoomorphology 83:1-47.

PAGE 147

136 Reif, W.-E. 1982. Evolution of dermal skel eton and dentition in vertebrates: the odonotode regulation theory. Evol utionary Biology 15:287-368. Reif, W.-E. 1983. Functional morphology a nd evolutionary ecology. Palontologie Zeitschrift 57:255-266. Reif, W.-E., D. McGill, and P. J. Motta. 1978. Tooth replacement rates of the sharks Triakis semifasciata and Ginglymostoma cirratum Zoologische Jahrbucher. Abteilung fur Anatomie und Ontogenie der Tiere 99:151-156. Richard, A. F., and R. E. Dewar. 1991. Lemur ecology. Annual Review of Ecology and Systematics 22:145-175. Sacco, T., and B. Van Valkenburgh. 2004. Ec omorphological indicators of feeding behaviour in the bears (Carnivora: Ursidae). Journal of Zoology, London 263:4154. Sanford, C. P. 2001. The novel "tounge-bite" apparatus" in the knifefish family Notopteridae (Teleostei: Osteoglossomorpha): are kinematic patterns conserved within a clade? Zoological Journa l of the Linnean Society 132:259-275. Sato, I., K. Shimada, and T. Sato. 1992. Morphology of the teeth of adult Caudata and Apoda: fine structure and chemistry of enamel. Journal of Morphology 214:341350. Sato, I., K. Shimada, A. Yokoi, J. C. Handal, N. Asuwa, and T. Ishii. 2005. Morphology of the teeth of the American alligator ( Alligator mississippiensis ): fine structure and chemistry of the enamel. Journal of Morphology 205:165-172.

PAGE 148

137 Schaeffer, B. 1967. Comments on elasmobranch evolution; pp. 3-35 i n P. W. Gilbert, R. F. Mathewson, and D. P. Rall (eds.), Sharks, Skates, and Rays. John Hopkins University Press, Baltimore. Schwenk, K. 2000. Tetrapod feeding in the co ntext of vertebrate morphology; pp. 3-20 i n K. Schwenk (ed.), Feeding: Form, Function, and Evolution in Tetrapod Vertebrates. Academic Press, New York. Schwimmer, D. R., J. D. Stewart, and G. D. Williams. 1997. Scavenging by sharks of the genus Squalicorax in the Late Cretaceous of North America. Palaios 12:71-83. Shimada, K., and G. E. Hooks III. 2004. Sharkbitten protostegid turtles from the Upper Cretaceous Mooreville Chalk, Alabam a. Journal of Paleontology 78:205-210. Shimizu, D., and G. A. Macho. 2007. Functional significance of the microstructural detail of the primate dentino-enamel junction: a possible example of exaptation. Journal of Human Evolution 52:103-111. Shirai, S. 1996. Phylogenetic interrelations hips of neoselachians (Chondricthyes: Euselachii); pp. 9-34 i n M. L. J. Stiassny, L. R. Parenti, and G. D. Johnson (eds.), Interrelationships of Fishes. Academic Press, San Diego. Shirai, S., and K. Nakaya. 1992. Functional morphology of feeding apparatus of the cookie-cutter shark, Isistius brasiliensis (Elasmobranchii, Da latiinae). Zoological Science 9:811-821. Simpfendorfer, C. A., A. B. Goodreid, and R. B. McAuley. 2001. Size, sex, and geographic variation in the diet of the tiger shark ( Galeocerdo cuvier ) in Western Australian waters. Environmen tal Biology of Fishes 61:37-46.

PAGE 149

138 Soler-Gijon, R. 1995. Evidence of predator-prey relationship in xenacanth sharks of the Upper Carboniferous (Stephanian C) from Puertollano, Spain. Geobios 28:151156. Springer, S. 1961. Dynamics of the feeding mechanisms in large galeoid sharks. American Zoologist 1:183-185. Springer, S. 1967. Social organiza tion of shark populations; pp. 149-176 i n P. W. Gilbert, R. F. Mathewson, and D. P. Rall (eds.), Sharks, Skates, and Rays. John Hopkins University Press, Baltimore. Stahl, B. J., and D. C. Parris. 2004. The complete dentition of Edaphodon mirificus (Chondrichthyes: Holocephali) from a single individual. Journal of Paleontology 78:388-392. Stavitzky, A. H. 1981. Hinged teeth in snakes : an adaptation for sw allowing hard-bodied prey. Science 212:346-349. Summers, A. P. 2000. Stiffening the stingray skeleton an investigation of durophagy in Myliobatid stingrays (Ch ondrichthyes, Batoidea, Myliobatidae). Journal of Morphology 243:113-126. Summers, A. P., and J. H. Long, Jr. 2006. The mechanical behavior of fish skeletal tissues; pp. 141-177 i n R. E. Shadwick and G. V. Lauder (eds.), Fish Biomechanics. Elsevier, New York. Summers, A. P., R. A. Ketcham, and T. Rowe. 2004. Structure and function of the horn shark ( Heterodontus francisci ) cranium through ontogeny: the development of a hard prey crusher. Journal of Morphology 260:1-12.

PAGE 150

139 Tricas, T. C., and J. E. McCosker. 1984. Predatory behavior of the white shark ( Carcharodon carcharias), with notes on its biology. Proceedings of the California Academy of Sciences 43:221-238. Turingan, R. G. 1994. Ecomorphological relati onships among Caribbean tetraodontiform fishes. Journal of Zoology, London 233:493-521. Turingan, R. G., P. C. Wainwri ght, and D. A. Hensley. 1995. Interpopulation variation in prey use and feeding biomechanics in Ca rribean triggerfishes. Oceologia 102:296304. Van Valkenburgh, B. 1988. Incidence of tooth breakage among large, predatory mammals. The American Naturalist 131:291-302. Van Valkenburgh, B. 1989. Carnivore dental ad aptations and diet: a study of trophic diversity within guilds; pp. 410-436 i n J. L. Gittleman (ed.), Carnivore Behavior, Ecology, and Evolution. Cornell University Press, New York. Van Valkenburgh, B., and C. B. Ruff. 1987. Ca nine tooth strength and killing behaviour in large carnivores. Jour nal of Zoology, London 212:379-397. Vogel, H. G. 1988. Age-dependent mechanical and biochemical changes in the skin. Bioengineering and the Skin 4:75-81. Vogel, S. 2003. Comparative Biomechanics: Life 's Physical World. Princeton University Press, Princeton, 580 pp. Wainwright, P. C. 1987. Biomechanical lim its to ecological performance: mollusccrushing by the Caribbean hogfish, Lachnolaimus maximus (Labridae). Journal of Zoology, London 213:283-297.

PAGE 151

140 Wainwright, P. C. 1988. Morphol ogy and ecology: func tional basis of feeding constraints in Caribbean labrid fishes. Ecology 69:635-645. Wainwright, P. C. 1994. Functi onal morphology as a tool in ecological research; pp. 4259 i n P. C. Wainwright and S. M. Re illy (eds.), Ecological Morhpology: Integrative Organismal Biology. Univer sity of Chicago Press, Chicago. Wainwright, P. C. 1996. Ecological explan ation through functional morphology: the feeding biology of sunfishes. Ecology 77:1336-1343. Wainwright, P. C., and B. A. Richard. 1995. Predicting patterns of prey use from morphology of fishes. Environmen tal Biology of Fishes 44:97-113. Waller, G. N. H., and A. Baranes. 1991. Chondrocranium morphology of norther Red Sea triakid sharks and relationships to feed ing habits. Journal of Fish Biology 38:715730. Wass, R. C. 1973. Size, growth, reproduction of the sandbar shark, Carcharhinus milberti in Hawaii. Pacific Science 27:305-318. Waters, N. E. 1980. Some mechanical and physi cal properties of teet h. Symposia of the Society for Experime ntal Biology 34:75-98. Wautier, K., C. Van der Heyen, and A. Huysseune. 2001. A quantitative analysis of pharyngeal tooth shape in the zebrafish ( Danio rerio, Teleostei, Cyprinidae). Archives of Oral Biology 46:67-75. Weins, J. A., and J. T. Rotenberry. 1980. Patterns of mor phology and ecology in grassland and shrubsteppe bird popula tions. Ecological M onographs 50:287-308.

PAGE 152

141 Whitenack, L. B., D. R. Elliott, and J. P. Brandenburg. 2002. A case study in paleoecology from the Mi ssissippian of Missouri, with a focus on chondricthyan icthyoliths. Transactions of the Missouri Academy of Sciences 36:11-14. Willemse, J. J. 1972. Arrangement of connective tissue fibres in the musculus lateralis of the spiny dogfish, Squalus acanthias L. (Chondrichthyes). Zoomorphology 72:231-244. Williams, E. E. 1972. The origin of faunas. E volution of lizard cogeners in a complex island fauna: a trial analysis. Evolutionary Biology 6:47-89. Williams, M. 2001. Tooth retention in cla dodont sharks: with a comparison between primitive grasping and swallowing, and modern cutting and gouging feeding mechanisms. Journal of Vert ebrate Paleontology 21:214-226. Williams, M. E. 1990. Feeding behavior in Cleveland Shale fishes; pp. 273-287 i n A. J. Boucot (ed.), Evolutionary Paleobiology of Behavior and Coevolution. Elsevier, Amsterdam. Williamson, T. E., J. I. Kirkland, and S. G. Lucas. 1993. Sleachians from the Greenhorn Cyclothem ("Middle" Cretaceous: Cenomanian-Turonian), Black Mesa, Arizona, and the paleogeographic dist ribution of Late Cretaceous selachians. Journal of Paleontology 67:447-474. Wroe, S., D. R. Huber, M. Lowry, C. R. McHenry, K. Moreno, P. D. Clausen, T. L. Ferrara, E. Cunningham, M. N. Dea n, and A. P. Summers. 2008. Threedimensional computer analysis of white shark jaw mechanics: how hard can a white shark bite? Jour nal of Zoology 2008:1-7.

PAGE 153

142 Yabe, H. 1998. Selachian fauna from the Upper Miocene Senhata Formation, Bosco Peninsula, Central Japan. Natural History Research, Special Issue 5:33-61. Yamakawa, K. 1959. Comparative-anatomical studi es on the enamel structure of rodents. Acta Anatomy Nipponica 34:852-866. Yamaoka, K., M. Hori, and S. Kuratani. 1986. Ecomorphology of feeding in 'goby-like' cichlid fishes in Lake Tanganyika Physiology & Ecology Japan 23:17-29. Zangerl, R. 1981. Chondrichthyes II. Handbook of Paleoicthyology. Gustav Fischer Verlag, New York, 114 pp. Zangerl, R., and E. S. Richardson, Jr. 1963. The paleocological history of two Pennsylvanian black shales. Fi eldiana: Geology Memoirs 4:1-352.

PAGE 154

143 Appendix A: Tooth Measurements Taken for Canonical Correlates Analysis

PAGE 155

Appendix A: See Chapter 1 for abbreviations. BCW, BW, CH, DCL, MCL, NCW, and NH are measured in millimeters. CA and NA are measured in degrees. 144 S. ringensH. gG. cC. leucasC. limbatusP. glaucaI. oxC. caS. mokarranN. bSpeciesI. ox AARCARBCWBWCHCADCLCIMean3.012.744.257.1510.830.0010.431.21SD0.881.171.350.610.760.000.290.11Mean1.321.094.407.235.350.004.601.65SD0.070.200.960.150.780.001.130.42Mean1.421.3616.4319.1022.200.0022.701.03SD0.070.073.332.863.430.003.370.04Mean0.880.8012.2212.729.480.008.581.73SD0.130.071.211.061.470.002.080.25Mean0.630.6621.2821.5613.2874.8014.801.38SD0.050.056.286.134.8912.876.550.21Mean0.850.7220.1822.1414.440.0015.021.34SD0.050.062.572.801.310.001.490.05Mean1.310.8212.6013.7510.250.0010.351.20SD0.190.071.272.190.210.000.070.01Mean0.930.8212.2813.969.980.0011.541.22SD0.090.052.132.691.270.001.730.53Mean1.670.9112.5314.3311.340.0012.161.05SD0.060.122.292.251.700.001.990.05Mean1.861.994.003.657.6097.607.800.98SD0.010.561.270.350.2815.840.710.05Mean0.590.3229.9030.839.7054.2017.731.18SD0.030.046.416.992.451.987.480.48riseusuvieryrinchus (l)rchariasrevirostrisyrinchus (a)

PAGE 156

Appendix A (continuted): 145 SpeciesI. oxI. oxC. caS. mokarranG. cC. leucasC. limbatusP. glaucaN. bS. ringensH. g MCLNCWNHNABOLATLCSEMean12.632.807.87148.97-14.56SD1.230.950.062.4914.21Mean7.352.803.70100.75-17.53SD0.070.140.0013.5112.04Mean23.3314.1019.90169.53-4.17SD3.023.013.478.698.33Mean14.507.506.60127.5515.67SD1.650.481.2512.1270.59Mean19.4810.606.60112.002.67SD5.903.701.9119.313.82Mean20.2211.359.68146.2321.58SD2.450.570.546.723.58Mean12.406.258.00125.703.60SD0.001.771.130.140.57Mean13.466.986.46116.6823.49SD4.760.981.073.753.83Mean12.705.409.00120.867.96SD2.131.101.674.095.41Mean7.604.408.20131.2029.29SD0.280.851.568.917.60Mean18.737.004.1583.800.00SD0.710.990.784.670.00yrinchus (a)yrinchus (l)rchariasuvierrevirostrisriseus020101010020201220121012012020002

PAGE 157

146 Appendix B: Finite Element Models

PAGE 158

Distal Mesial Labial Lingual Distal Mesial Labial Lingual161Appendix B B1: Hexanchus griseus, distal draw 147

PAGE 159

162Appendix B (Continued) Distal Mesial Labial Lingual B2: Hexanchus griseus, labial hold 148

PAGE 160

163Appendix B (Continued) Distal Mesial Labial Lingual B3: Hexanchus griseus, lingual hold 149

PAGE 161

164Appendix B (Continued) Distal Mesial Labial Lingual B4: Hexanchus griseus, mesial draw 150

PAGE 162

165Appendix B (Continued) Distal Mesial Labial Lingual B5: Hexanchus griseus, notch 151

PAGE 163

166Appendix B (Continued) Distal Mesial Labial Lingual B6: Hexanchus griseus, puncture 152

PAGE 164

167Appendix B (Continued) Distal Mesial Labial Lingual B7: Prionace glauca distal draw 153

PAGE 165

168Appendix B (Continued) Distal Mesial Labial Lingual B8: Prionace glauca labial hold 154

PAGE 166

169Appendix B (Continued) Distal Mesial Labial Lingual B9: Prionace glauca lingual hold 155

PAGE 167

170Appendix B (Continued) Distal Mesial Labial Lingual B10: Prionace glauca mesial draw 156

PAGE 168

171Appendix B (Continued) Distal Mesial Labial Lingual B11 : Prionace glauca notch 157

PAGE 169

172Appendix B (Continued) Distal Mesial Labial Lingual B12: Prionace glauca puncture 158

PAGE 170

173Appendix B (Continued) Distal Mesial Labial Lingual B13: Carcharhinus leucas, distal draw 159

PAGE 171

174Appendix B (Continued) Distal Mesial Labial Lingual B14: Carcharhinus leucas, labial hold 160

PAGE 172

175Appendix B (Continued) Distal Mesial Labial Lingual B15: Carcharhinus leucas, distal draw 161

PAGE 173

176Appendix B (Continued) Distal Mesial Labial Lingual B16: Carcharhinus leucas, mesial draw 162

PAGE 174

177Appendix B (Continued) Distal Mesial Labial Lingual B18: Carcharhinus leucas, notch 163

PAGE 175

178Appendix B (Continued) Distal Mesial Labial Lingual B18: Carcharhinus leucas, puncture 164

PAGE 176

179Appendix B (Continued) Distal Mesial Labial Lingual B19: Cladodus sp., distal draw 165

PAGE 177

180Appendix B (Continued) Distal Mesial Labial Lingual B20: Cladodus sp., labial hold 166

PAGE 178

182Appendix B (Continued) Distal Mesial Labial Lingual B22: Cladodus sp., mesial draw 168

PAGE 179

183Appendix B (Continued) Distal Mesial Labial Lingual B23: Cladodus sp., puncture 169

PAGE 180

184Appendix B (Continued) Distal Mesial Labial Lingual B24: Carcharhinus limbatus, distal draw 170

PAGE 181

185Appendix B (Continued) Distal Mesial Labial Lingual B25: Carcharhinus limbatus, labial hold 171

PAGE 182

186Appendix B (Continued) Distal Mesial Labial Lingual B26: Carcharhinus limbatus, lingual hold 172

PAGE 183

187Appendix B (Continued) Distal Mesial Labial Lingual B27: Carcharhinus limbatus, mesial draw 173

PAGE 184

188Appendix B (Continued) Distal Mesial Labial Lingual B28: Carcharhinus limbatus, puncture 174

PAGE 185

189Appendix B (Continued) Distal Mesial Labial Lingual B29: Sphyrna mokarran, distal draw 175

PAGE 186

190Appendix B (Continued) Distal Mesial Labial Lingual B30: Sphyrna mokarran, labial hold 176

PAGE 187

191Appendix B (Continued) Distal Mesial Labial Lingual B31: Sphyrna mokarran, lingual hold 177

PAGE 188

192Appendix B (Continued) Distal Mesial Labial Lingual B32: Sphyrna mokarran, mesial draw 178

PAGE 189

193Appendix B (Continued) Distal Mesial Labial Lingual B33: Sphyrna mokarran, puncture 179

PAGE 190

194Appendix B (Continued) Distal Mesial Labial Lingual B34: Sphyrna mokarran, notch 180

PAGE 191

195Appendix B (Continued) Distal Mesial Labial Lingual B35: Hybodus sp., distal draw 181

PAGE 192

196Appendix B (Continued) Distal Mesial Labial Lingual B36: Hybodus sp., labial hold 182

PAGE 193

197Appendix B (Continued) Distal Mesial Labial Lingual B37: Hybodus sp., lingual hold 183

PAGE 194

198Appendix B (Continued) Distal Mesial Labial Lingual B38: Hybodus sp., mesial draw 184

PAGE 195

199Appendix B (Continued) Distal Mesial Labial Lingual B39: Hybodus sp., puncture 185

PAGE 196

200Appendix B (Continued) Distal Mesial Labial Lingual B40: Negaprion brevirostris distal draw 186

PAGE 197

201Appendix B (Continued) Distal Mesial Labial Lingual B41: Negaprion brevirostris labial hold 187

PAGE 198

202Appendix B (Continued) Distal Mesial Labial Lingual B42: Negaprion brevirostris lingual hold 188

PAGE 199

203Appendix B (Continued) Distal Mesial Labial Lingual B43: Negaprion brevirostris mesial draw 189

PAGE 200

204Appendix B (Continued) Distal Mesial Labial Lingual B44: Negaprion brevirostris puncture 190

PAGE 201

205Appendix B (Continued) Distal Mesial Labial Lingual B45: Isurus oxyrinchus, anterior tooth, distal draw 191

PAGE 202

206Appendix B (Continued) Distal Mesial Labial Lingual B46: Isurus oxyrinchus, anterior tooth, labial hold 192

PAGE 203

207Appendix B (Continued) Distal Mesial Labial Lingual B47: Isurus oxyrinchus, anterior tooth, lingual hold 193

PAGE 204

208Appendix B (Continued) Distal Mesial Labial Lingual B48: Isurus oxyrinchus, anterior tooth, mesial draw 194

PAGE 205

209Appendix B (Continued) Distal Mesial Labial Lingual B49: Isurus oxyrinchus, anterior tooth, puncture 195

PAGE 206

210Appendix B (Continued) Distal Mesial Labial Lingual B50: Isurus oxyrinchus, lateral tooth, distal draw 196

PAGE 207

211Appendix B (Continued) Distal Mesial Labial Lingual B51: Isurus oxyrinchus, lateral tooth, labial hold 197

PAGE 208

212Appendix B (Continued) Distal Mesial Labial Lingual B52: Isurus oxyrinchus, lateral tooth, lingual hold 198

PAGE 209

213Appendix B (Continued) Distal Mesial Labial Lingual B53: Isurus oxyrinchus, lateral tooth, mesial draw 199

PAGE 210

214Appendix B (Continued) Distal Mesial Labial Lingual B54: Isurus oxyrinchus, lateral tooth, puncture 200

PAGE 211

215Appendix B (Continued) Distal Mesial Labial Lingual B55: Scymnodon ringens, distal draw 201

PAGE 212

216Appendix B (Continued) Distal Mesial Labial Lingual B56: Scymnodon ringens, labial hold 202

PAGE 213

217Appendix B (Continued) Distal Mesial Labial Lingual B57: Scymnodon ringens, lingual hold 203

PAGE 214

218Appendix B (Continued) Distal Mesial Labial Lingual B58: Scymnodon ringens, mesial draw 204

PAGE 215

219Appendix B (Continued) Distal Mesial Labial Lingual B59: Scymnodon ringens, puncture 205

PAGE 216

220Appendix B (Continued) Distal Mesial Labial Lingual B60: Galeocerdo cuvier distal draw 206

PAGE 217

221Appendix B (Continued) Distal Mesial Labial Lingual B61: Galeocerdo cuvier labial hold 207

PAGE 218

222Appendix B (Continued) Distal Mesial Labial Lingual B62: Galeocerdo cuvier lingual hold 208

PAGE 219

223Appendix B (Continued) Distal Mesial Labial Lingual B63: Galeocerdo cuvier mesial draw 209

PAGE 220

224Appendix B (Continued) Distal Mesial Labial Lingual B64: Galeocerdo cuvier notch 210

PAGE 221

225Appendix B (Continued) Distal Mesial Labial Lingual B65: Galeocerdo cuvier puncture 211

PAGE 222

226Appendix B (Continued) Distal Mesial Labial Lingual B66: Carcharodon carcharias, distal draw 212

PAGE 223

227Appendix B (Continued) Distal Mesial Labial Lingual B67: Carcharodon carcharias, labial hold 213

PAGE 224

228Appendix B (Continued) Distal Mesial Labial Lingual B68: Carcharodon carcharias, lingual hold 214

PAGE 225

229Appendix B (Continued) Distal Mesial Labial Lingual B69: Carcharodon carcharias, mesial draw 215

PAGE 226

230Appendix B (Continued) Distal Mesial Labial Lingual B70: Carcharodon carcharias, puncture 216

PAGE 227

231 231Appendix B (Continued) Side Labial Lingual Loaded Side B71: Xenacanthus compressus, draw 217

PAGE 228

232 232Appendix B (Continued) Side Labial Lingual B72: Xenacanthus compressus, labial hold 218

PAGE 229

233 233Appendix B (Continued) Side Labial Lingual B73: Xenacanthus compressus, lingual hold 219

PAGE 230

220 Appendix C: Specimens Used for Evolutionary Analysis

PAGE 231

Appendix C: AMNH: American Museum of Natural History, FLMNH = Florida Museum of Natural History, FMNH = Field Museum of Natural History, GH = Private collection of Gordon Hubbell, NMNH = National Museum of Natural History (Smithsonian Institution), USF = University of South Florida, UT = University of Tampa. 221HetOrectolobidaeHemiscyllidaeStegostomGinglymMit FamilySpeciesCollectionSpecimenTL (cm)SexerodontidaeHeterodontus francisciGHWALL86FUTA74MUTB72FUTC67MUTDES 06-0969FOrectolobus maculatusFMNH86308--GH---GH---NMNH3999965FNMNH4000452FChiloscyllium plagiosumGH-58FatidaeStegostoma fasciatumGHBIN--GH"Phuket, Thailand"--ostomidaeGinglymostoma cirratumFLMNH48301228-FLMNH209077244FGH1-27-99243MGH"207 lb."248MGH30-3-99245MsukurinidaeMitsukurina owstoniNMNH50972-F

PAGE 232

Appendix C (continued): 222AloOdoPLa piidaeAlopias superciliosusFLMNH010302002.01--NMNH110927--USFASUP01244-USFASUP02--ntaspididaeCarcharias taurusUSFCTAUR01--FLMNH47900--FLMNH19705007.017240-FMNH31193--seudocarcharinidaePseudocarcharias kamoharaiFLMNH47481102FFLMNH14775886FFLMNHBURGESS--FMNH117471101MNMNH309254107MmnidaeIsurus oxyrinchusUSFLISAMAKO42305229-FLMNH30102013.28163FFLMNHISHAF--GHISUR-1-18239MUSFISUR02--Carcharias charcharodonAMNH53095--FLMNH48285262FFLMNH20105018.01237MNMNH27374--NMNH110889-

PAGE 233

Appendix C (continued): 223ScPTriaCa Lamna nasusNMNH125884--FMNH51197--USFLNAS01218MyliorhinidaeScyliorhinus retiferFLMNH3635934MFLMNH3635938MFLMNH3635940MFLMNH3635932FGaleus araeGH"Cape Canaveral"28FParamaturus xaniurusFLMNH16678248FseudotriakidaePseudotriakis microdonGH"Senegal 4-13-98"216MkidaeTriakis semifasciataGHCASE158FrcharhinidaeCarcharhinus leucasFLMNH39601009.01278FFLMNH20229229FUSFCLEU05244-USFCLEU02226-USFCLEU04340-Carcharhinus limbatusFLMNH110301939.019178FFLMNH40501914.03156FGaleocerdo cuvierFLMNHDES 06-09350MFLMNH30501903.03332MFLMNH50501903.83228FFLMNH70301302.02219MFLMNH79801029327F

PAGE 234

Appendix C (continued): 224Sphy Carcharhinus pereziFLMNH208612--FLMNH209074224FFLMNH209076--FLMNH604012016.9190MUSFCPERE03--Carcharhinus plumbeusUSFCPLU07191FUSFCPLU08180FUSFCPLU06178FNegaprion brevirostrisFLMNHZ9239--FLMNH208854213-FLMNH144750--FLMNH208425--FMNH51202--Prionace glaucaAMNH89126237MAMNH42154--AMNH89229USFPRIO01290-USFPRIO02-MrinidaeSphyrna mokarranFLMNH030101020.16350-FLMNH040401902.18319FFLMNH079901007.04300MFLMNH080501902.121305MFLMNH110405937.20347F

PAGE 235

Appendix C (continued): 225HeC Sphyrna tiburoUSFLISATIB-FUSFSTIB0289FUSFSTIB0392FUSFSTIB0488FUSFSTIB0684FmigaleidaeHemipristis elongataAMNH89025130MAMNH8902697MAMNH8903893FFLMNH48116--NMNH263799--Paragaleus pectoralisAMNH79928108FNMNH23297998FNMNH232981104MNMNH232980113MHemigaleus microstomaAMNH8903686MAMNH8903761FAMNH8904080MAMNH8904170FhlamydoselachidaeChlamydoselachus anguineusFLMNH1665126MFMNH34236--GH8-26-98162FNMNH48530-MNMNH203466-M

PAGE 236

Appendix C (continued): 226HexanchidaNotoryEchinorhinidaeSomEt eHexanchus griseusAMNH78171252FAMNH78173294FNMNH104474108MNMNH188048433FUSFHEXG01297FnchidaeNotorynchus cepedianusGH---GH"Humbolt"246FGHWALL225MNMNH8768166MEchinorhinus cookeiFLMNH103000244FGH2-8-1999213-GH"3-4-01 Henke"208FGH"Near Ventura"203FniosidaeScymnodon ringensGHWALL96FUSFSCYM01--Somniosus microcephalusUSFSOMN01305FAMNH78352--FMNH51405--mopteridaeAculeola nigraNMNH22020844MNMNH22020842MEtmopterus virensFLMNH2795021MFLMNH2795023FFLMNH2795021FFLMNH14824626F

PAGE 237

Appendix C (continued): 227OxDaCentrophorSquSquPristiophor Etmopterus luciferFLMNH4166845FFLMNH4166847FFLMNH4166834FynotidaeOxynotus bruniensisGHCASE67FlatiidaeDalatias lichaAMNH78306110MAMNH78308116MFMNH33944--USFDALA01--USFDALA02--idaeCentrophorus granulosusFLMNH3016085MFLMNH3016388FFLMNH7957994MFLMNH161527104FFLMNH161527107FalidaeSqualus acanthiasUSFSACAN0132FSACAN0226FSACAN0325FSACAN0431FSACAN0528FatidaeSquatina dumerilGH12-9-1999125MNMNH110892--idaePristiophorus cirratumGHCASE137

PAGE 238

Appendix D: Dietary References for Evolutionary Analysis 228

PAGE 239

Appendix D: Alonso, M. K., E. A. Crespo, N. A. Garcia, S. N. Pedraza, P. A. Mariotti, and N. J. Mora. 2002. Fishery and ontogenetic changes in the diet of the spiny dogfish, Squalus acanthias, in Patagonian waters, Argentina. Environmental Biology of Fishes 63:193-202. Avsar, D. 2001. Age, growth, reproduction, and feeding of the spurdog (Squalus acanthias Linnaeus, 1758) in the south-eastern Black Sea. Estuarine Costal and Shelf Science 52:269-278. Baba, O., T. Taniuchi, and Y. Nose. 1987. Depth distribution and food habits of three species of small squaloid sharks off Choshi. Nippon Suisan Gakkaishi 53:417-424. Balart, E. F., H. Gonzalez-Garcia, and C. Villevenico-Garayzar. 2000. Notes on the biology of Cephalurus cephalus and Paramaturus xaniurus (Chondrichthyes: Scyliorhinidae) from the west coast of Baja, California Sur, Mexico. Fisheries Bulletin 98:219-221. Bethea, D. M., J. A. Buckel, and J. K. Carlson. 2004. Foraging ecology of the early life stages of four sympatric shark species. Marine Ecology Progress Series 268:245-264. Bethea, D. M., L. Hale, J. K. Carlson, E. Cortes, C. A. Manire, and J. Gelsleichter. 2007. Latitudinal varation in the diet and daily ration of the bonnethead shark, Sphyrna tiburo, from the eastern Gulf of Mexico. Marine Biology 152:359-363. 229

PAGE 240

Appendix D (continued): Castro, J. I. 2000. The biology of the nurse shark, Ginglymostoma cirratum, off the Florida east coast and the Bahama Islands. Environmental Biology of Fishes 58:1-22. Castro, J. I., P. M. Bubcucis, and N. A. Overstrom. 1998. The reproductive biology of the chain dogfish, Scyliorhinus retifer. Copeia 1998:740-746. Cherel, Y., and G. Duhamel. 2004. Antarctic jaws: cepahlopod prey of sharks in Kerguelen waters. Deep-Sea Research I 51:17-31. Clark, M. R., D. C. Clark, H. R. Martins, and H. R. DaSilva. 1996. The diet of the blue shark (Prionace glauca) in Azorean waters. Arquipelago: Life and Marine Science 14A:41-56. Cliff, G. 1995. Sharks caught in the protective gill nets off Kwazulu-Natal, South Africa. 8. The great hammerhead shark Sphyrna mokarran (Ruppell). South African Journal of Marine Science 15:105-114. Cliff, G., and R. Dudley. 1991. Sharks cught in the protective gill nets off Natal, South Africa. 4. The bull shark Carcharhinus leucas Valenciennes. South African Journal of Marine Science 10:253-279. Collard, S. B. 1970. Forage of some eastern Pacific midwater fishes. Copeia 1970:348-354. Compagno, L. J. V. 1984a. FAO Species Catalogue. Sharks of the World. An annotated and illustrated catalogue of shark species known to date, Part 1: Hexanchiformes to lamniformes. Food and Agriculture Organization of the United Nations, Rome. 230

PAGE 241

Appendix D (continued) Compagno, L. J. V. 1984b. FAO Species Catalogue. Sharks of the World. An annotated and illustrated catalogue of shark species known to date, Part 2: Carcharhiniformes. Food and Agriculture Organization of the United Nations, Rome. Compagno, L. J. V. 2001. Sharks of the World: An annotated and illustrated catalogue of shark species known to date. Volume 2. Bullhead, mackerel and carpet sharks (Heterodontiformes, Lamniformes and Orectolobiformes). FAO Species Catalogue for Fishery Purposes. 2:1-269. Compagno, L. J. V., M. Dando, and S. Fowler. 2005. Sharks of the World. Princeton University Press, Princeton, 480 pp. Cortes, E. 1999. Standardized diet compositions and trophic levels of sharks. ICES Journal of Marine Science 56:707-717. Cortes, E., C. A. Manire, and R. E. Hueter. 1996. Diet, feeding habits, and diel feeding chronology of the bonnethead shark Sphyrna tiburo in southwest Florida. Bulletin of Marine Science 58:353-367. Crespi-Abril, A. C., N. A. Garcia, E. A. Crespo, and M. A. Coscaralla. 2003. Consumption of marine mammals by broadnose sevengill shark Notorhynchus cepedianus in the northern and central Patagonian Shelf. Latin American Journal of Aquatic Mammals 2:101-107. 231

PAGE 242

Appendix D (continued): Cross, J. N. 1988. Aspects of the biology of two scyliorhinid sharks, Apristurus brunneus and Paramaturus xaniurus, from the upper continental slope off southern California. U.S. Fish and WIldlife Service Fishery Bulletin 86:691-702. Demirhan, S. A., and K. Seyhan. 2007. Life history of the spiny dogfish, Squalus acanthias (L. 1758), in the southern Black Sea. Fisheries Research 85:210-216. DiBeneditto, A. P. M. 2004. Presence of franciscana dolphin (Pontoporia blainvillei) remains in the stomach of a tiger shark (Galeocerdo cuvier) captured in southeastern Brazil. Aquatic Mammals 30:311-314. Dowd, W. W., R. W. Brill, P. G. Bushnell, and J. A. Musick. 2006. Estimating consumption rates of juvenile sandbar shark (Carcharhinus plumbeus) in Chesapeake Bay, Virginia, using a bioenergetic model. Fishery Bulletin 104:332-342. Dudley, S. F. J., and G. Cliff. 1993. Sharks caught in the protective gill nets off Natal, South Africa. 7. The blacktip shark Carcharhinus limbatus (Valenciennes). South African Journal of Marine Science 13:237-254. Duffy, C. 1997. Further records of the goblin shark, Mitsukurina owstoni (Lamniformes: Mitsukurinidae) from New Zealand. New Zealand Journal of Zoology 24:167-171. Duffy, K. A. 1999. Feeding, growth, and bioenergetics of the chain dogfish, Scyliorhinus retifer. Oceanography, University of Rhode Island, Kingston, 177 pp. 232

PAGE 243

Appendix D (continued): Ebert, D. A. 1994. Diet of the sixgill shark Hexanchus griseus off southern Africa. South African Journal of Marine Science 14:213-218. Ebert, D. A. 2002. Ontogenetic changes in the diet of the sevengill shark (Notorhynchus cepedianus). Marine and Freshwater Research 53:517-523. Ebert, D. A., and T. B. Ebert. 2005. Reproduction, diet and habitat use of leopard sharks, Triakis semifasciata (Girard), in Humbolt Bay, California, USA. Marine and Freshwater Research 56:1089-1098. Garibaldi, F., and L. O. Relini. 2000. Summer abundance, size, and feeding habits of the blue shark, Prionace glauca, in the pelagic sanctuary of the Ligurian Sea. Biologia Marina Mediterranea 7:324-333. Gelsleichter, J., J. A. Musick, and S. Nichols. 1999. Food habits of the smooth dogfish, Mustelus canis, dusky shark, Carcharhinus obscurus, Atlantic sharpnose shark, Rhizoprionidon terraenovae, and the sand tiger, Carcharias taurus, from the northwest Atlantic Ocean. Environmental Biology of Fishes 54:205-217. Golani, D., and S. Pisanty. 2000. Biological aspects of the gulper shark, Centrophorus granulosus (Blotch and Schneider, 1801) from the Mediterranean coast of Israel. Journal of the Marine Biological Association of the United Kingdom 88:411-414. Hanchet, S. 1991. Diet of spiny dogfish, Squalus acanthias, on the east coast, South Island, New Zealand. Journal of Fish Biology 39:313-323. 233

PAGE 244

Appendix D (continued): Heithaus, M. R. 2001. The biology of tiger sharks (Galeocerdo cuvier) in Shark Bay, Western Australia: sex ratio, size distribution, diet, and seasonal changes in catch rates. Environmental Biology of Fishes 61:25-36. Henderson, A. C., K. Flannery, and J. Dunne. 2001. Observations on the biology and ecology of the blue shark in the north-east Atlantic. Journal of Fish Biology 58:1346-1358. Henderson, A. C., J. Dunne, and K. Flannery. 2002. Stomach contents of spiny dogfish, Squalus acanthias L. off the west coast of Ireland. Irish Naturalists' Journal 27:101-105. Henderson, A. C., K. Flannery, and J. Dunne. 2003. Biological observations on shark species taken in commericial fisheries to the west of Ireland. Biology and Environment 103B:1-7. Hoffmayer, E. R., and G. R. Parsons. 2003. Food habits of three shark species from the Mississsippi Sound in the northern Gulf of Mexico. Southeast Naturalist 2:271-280. Huveneers, C., N. M. Otway, S. E. Gibbs, and R. G. Harcourt. 2007. Quantitative diet assessment of wobegong sharks (genus Orectolobus) in New South Wales, Australia. ICES Journal of Marine Science 64:1272-1281. Jones, B. C., and G. H. Geen. 1977. Food and feeding of spiny dogfish (Squalus acanthias) in British Columbia waters. Journal of the Fisheries Research Board of Canada 34:2067-2078. 234

PAGE 245

Appendix D (continued): Joyce, W. N., S. E. Campana, L. J. Natanson, N. E. Kohler, H. L. Pratt, Jr., and C. F. Jensen. 2002. Analysis of stomach contents of the porbeagle shark (Lamna nasus Bonnaterre) in the northwest Atlantic. Journal of Marine Science 59:1263-1269. Kabasakal, H. 2004. Preliminary observations on the reproductive biology and diet of the bluntnose sixgill shark, Hexanchus griseus (Bonnaterre, 1788) (Chondrichthyes: Hexanchidae) in Turkish seas. Acta Adriatica 45:187-196. Kabasakal, H., and E. Kabasakal. 2002. Morphometrics of young kitefin sharks, Dalatias licha (Bonaterre, 1788), from northeastern Aegean Sea, with notes on its biology. Annales Anali za Istrske in Mediteranske Studike 12:161-166. Kubodera, T., H. Watanabe, and T. Ichii. 2007. Feeding habits of the blue shark, Prionace glauca, and salmon shark, Lamna ditropsis, in the transition region of the western north Pacific. Reviews in Fish Biology and Fisheries 17:111-124. Kubota, T., Y. Shiobara, and T. Kubodera. 1991. Food habits of the frilled shark Chlamydoselachius anguineus collected from Surugua Bay, Central Japan. Nippon Suisan Gakkaishi 57:15-20. Laptikhovshy, V. V., A. I. Arkhipkin, and A. C. Henderson. 2001. Feeding habits and dietary overlap in spiny dogfish Squalus acanthias (Squalidae) and narrowmouth catshark Schroederichthys bivius (Scyliorhinidae). Journal of the Marine Biological Association of the United Kingdom 81:1015-1018. Lessa, R. P., and A. Almeida. 1998. Feeding habits of the bonnethead shark, Sphyrna tiburo, from northern Brazil. Cybium 22:383-394. 235

PAGE 246

Appendix D (continued): Lowe, C. G., B. M. Wetherbee, G. L. Crow, and A. L. Tester. 1996. Ontogenetic dietary shifts and feeding behavior of the tiger shark, Galeocerdo cuvier, in Hawaiian waters. Environmental Biology of Fishes 47:203-211. Lowry, D., and P. J. Motta. 2007. Ontogeny of feeding behavior and cranial morphology in the whitespotted bamboo shark Chiloscylium plagiosum. Marine Biology 151:2013-2023. Lucifora, L. O., R. C. Menni, and A. H. Escalante. 2005. Reproduction, abundance, and feeding habits of the broadnose sevengill shark, Notorhynchus cepedianus, in north Patagonia, Argentina. Marine Ecology Progress Series 289:237-244. Lucifora, L. O., V. B. Garcia, R. C. Menni, and A. H. Escalante. 2006. Food habits, selectivity, and foraging modes of the schools shark, Galeorhinus galeus. Marine Ecology Progress Series 315:259-270. Maia, A., N. Quieroz, J. P. Correia, and H. Cabral. 2006. Food habits of the shortfin mako, Isurus oxyrinchus, off the southwest coast of Portugal. Environmental Biology of Fishes 77:157-167. Martin, R. A., N. Hammerschlag, R. S. Collier, and C. Fallows. 2005. Predatory behavior of white sharks (Carcharodon carcharias) at Seal Island, South Africa. Journal of the Marine Biological Association of the United Kingdom 85:1121-1135. McCord, M. E., and S. E. Campana. 2003. A quantitative assessment of the diet of the blue shark (Prionace glauca) off Nova Scotia, Canada. Journal of Northwest Atlantic Fisheries Science 32:57-63. 236

PAGE 247

Appendix D (continued): McElroy, W. D., B. M. Wetherbee, C. S. Mostello, C. G. Lowe, G. L. Crow, and R. C. Wass. 2006. Food habits and ontogenetic changes in diet of the sandbar shark, Carcharhinus plumbeus, in Hawaii. Environmental Biology of Fishes 76:81-92. Medved, R. J., C. E. Stillwell, and J. J. Casey. 1985. Stomach contents of young sandbar sharks Carcharhinus plumbeus, in Chincoteague Bay, Virginia. U.S. Fish and WIldlife Service Fishery Bulletin 83:395-402. Megalafonou, P., and A. Chatzispyrou. 2006. Sexual maturity and feeding of the gulper shark, Centrophorus granulosus, from the Eastern Mediterranean Sea. Cybium 30:67-74. Morato, T., E. Sola, M. P. Gros, and G. Mendezes. 2003. Diets of thornback rays (Raja clavata) and tope shark (Galeorhinus galeus) in the bottom longline fishery of the Azores, Northeastern Atlantic. Fisheries Bulletin 101:590-602. Papastamatiou, Y. P., B. M. Wetherbee, C. G. Lowe, and G. L. Crow. 2006. Distribution and diet of four species of carcharhinid shark in the Hawaiian Islands: evidence for resource partitioning and competitive exclusion. Marine Ecology Progress Series 320:239-251. Polo-Silva, C., A. Baigorri-Santacruz, F. Galvan-Magana, M. Grijalba-Bendeck, and A. Sanjuan-Munoz. 2007. Habitos alimentarios del tiburon zorro Alopias superciliosus (Lowe, 1839), en el Pacifico ecuatoriano. Revista de Biologia Marina y Oceanogradia 42:59-69. 237

PAGE 248

Appendix D (continued): Salini, J. P., S. J. M. Balaber, and D. T. Brewer. 1992. Diets of sharks from estuaries and adjacent waters of the north-eastern Gulf of Carpentaria, Australia. Australian Journal of Marine and Freshwater Research 43:87-96. Schmidt, T. W. 1985. Food of young juvenile lemon sharks, Negaprion brevirostris (Poey), near Sandy Key, western Florida Bay. Florida Scientist 49:7-10. Schwartz, F. J. 2000. Food of tiger sharks Galeocerdo cuvier (Carcharhinidae) from the northwest Atlantic Ocean, off North Carolina. Journal of the Elisha Mitchell Scientific Society 116:351-355. Segura-Zarzosa, J. C., L. A. Abitia-Cardenas, and F. Galvan-Magana. 1997. Observations on the feeding habits of the shark Heterodontus francisci Girard 1854 (Chondrichthyes: Heterodontidae) in San Ignacio Lagoon, Baja California Sur, Mexico. Ciencias Marinas 23:111-128. Simpfendorfer, C. A. 1992. Biology of tiger sharks (Galeocerdo cuvier) caught by the Queensland Shark Meshing Program off Townsville, Australia. Australian Journal of Marine and Freshwater Research 43:33-43. Simpfendorfer, C. A., A. B. Goodreid, and R. B. McAuley. 2001. Size, sex, and geographic variation in the diet of the tiger shark (Galeocerdo cuvier) in Western Australian waters. Environmental Biology of Fishes 61:37-46. Smale, M. J. 2005. The diet of the ragged-tooth shark (Carcharias taurus) Rafinesque 1810 in the Eastern Cape, South Africa. African Journal of Marine Science 27:331-335. 238

PAGE 249

Appendix D (continued): Smale, M. J., and G. Cliff. 1998. Cephalopods in the diets of four shark species (Galeocerdo cuvier, Sphyrna lewini, S. zygaena, and S. mokarran) from Kwazulu-Natal, South Africa. South African Journal of Marine Science 20:241-253. Snelson, F. F., T. J. Mulligan, and S. E. Williams. 1984. Food habits, occurrence, and population structure of the bull shark, Carcharhinus leucas, in Florida costal lagoons. Bulletin of Marine Science 34:71-80. Soto, J. M. R. 2001. On the presence of the caribbean reef shark, Carcharhinus perezi (Poey, 1876) (Chondrichthyes, Carcharhinidae), in the southwest Atlantic. Mare Magnum 1:135-139. Stevens, J. D., and G. L. Cuthbert. 1983. Observations on the identification and biology of Hemigaleus (Sleachii: Carcharhinidae) from Australian waters. Copeia 1983:487-497. Stevens, J. D., and J. M. Lyle. 1989. Biology of three hammerhead sharks (Eusphyrna blochii, Sphyrna mokarran, and S. lewini) from northern Australia. Australian Journal of Marine and Freshwater Research 40:129-146. Stewart, A. 2001. First record of the crocodile shark Pseudocarcharias kamoharai (Chondrichthyes: Lamniformes) from New Zealand waters. New Zealand Journal of Marine and Freshwater Research 35:1001-1006. Stillwell, C. E., and N. E. Kohler. 1982. Food, feeding habits, and estimates of daily ration of the shortfin mako (Isurus oxyrinchus) in the northwest Atlantic. Canadian Journal of Fisheries and Aquatic Science 39:407-414. 239

PAGE 250

Appendix D (continued): Stillwell, C. E., and N. E. Kohler. 1993. Food habits of the sandbar shark Carcharhinus plumbeus off the U.S. Northeast coast, with estimates of daily ration. U.S. Fish and WIldlife Service Fishery Bulletin 91:138-150. Tavares, R., and F. Provenzo. 2000. Feeding habits of the blacktip shark juveniles, Carcharhinus limbatus (Balenciennes, 1839) at the Archipelago Los Roques National Park, Venezuela. Acta Biologica Venezuela 20:59-67. Yano, K., and J. A. Musick. 1992. Comparison of morphometrics and Pacific specimens of the false catshark, Pseudotriakis microdon, with notes on stomach content. Copeia 1992:877-886. Yano, K., J. D. Stevens, and L. J. V. Compagno. 2007. Distribution, reproduction, and feeding of the Greenland shark Somniosus (Somniosus) microcephalus, with notes on two other sleeper sharks, Somniosus (Somniosus) pacificusi and Somniosus (Somniosus) antarcticus. Journal of Fish Biology 70:374-390. 240

PAGE 251

Appendix E: Discrete Variables for Evolutionary Analysis 241

PAGE 252

Appendix E: See Chapter 3 for abbreviations. 242 SpHeOrectolobus mGiStegostChMitsuPseudoCaAlLamIsuCaScGaPaPseudoTrHeHePaSpSp eciesDepthHabitatElasmoTeleostShrimpCrabCephWormEchMolMammalterodontus francisci0000111010aculatus0311011000nglymostoma cirratum0001011000oma fasciatum0001110010iloscyllium plagiosum0001010100kurina owstoni1200001000carcharias kamoharai2101101000rcharias taurus1011000000opias superciliosus2401001000na nasus2411001000rus oxyrhincus2401001000rcharias charcharodon1411000001yliorhinus retifer0301101000leus arae0300100000ramaturus xaniurus0300110000triakis microdon1201000000iakis semifasciata1001110100mipristis elongata1001001000migaleus microstoma1000001000ragaleus pectoralis1000001000hyrna mokarran1411000000hyrna tiburo1000110000

PAGE 253

Appendix E (continued): See Chapter 3 for abbreviations. 243 SpeciesHeterodontus francisciOrectolobus mGiStegostomChMitsuPseudocarcharias kamCaAlLamIsurus oxyrhinCarcharias chaScyliorhGaleus araeParamPseudotriakis mTriakis semHemHemParagaleus pectoralisSphyrna mSphyrna t SoftMedHardSE LowerSE UpperLC LowerLC UpperLATMT101001111aculatus101000000nglymostoma cirratum101001110a fasciatum011001110iloscyllium plagiosum111002220kurina owstoni100002200oharai110002100rcharias taurus010001210opias superciliosus110001200na nasus110001210cus110002200rcharodon010221100inus retifer110001110000001110aturus xaniurus00100??10icrodon010001111ifasciata111001110ipristis elongata110121000igaleus microstoma100021100100021101okarran010220100iburo001021101

PAGE 254

Appendix E (continued): See Chapter 3 for abbreviations. 244 SpCaCaCaCaNegaprion bPriGaCentSqSqPrOxynotus bruniScymSoDaAculEtEtEchiHeNoCh eciesDepthHabitatElasmoTeleostShrimpCrabCephWormEchMolMammalrcharhinus perezi1001000000rcharhinus leucas1011000001rcharhinus plumbeus1001011000rcharhinus limbatus1001000000revirostris1011000000onace glauca2101001000leocerdo cuvier0411010000rophorus granulosus1201001000ualus acanthias1001101000uatina dumeril0001000000istiophorus cirratum1301110000ensis02????????nodon ringens1201110000mniosus microcephalus1301000001latias licha1211000000eola nigra1301111000mopterus virens1200001000mopterus lucifer1201001000norhinus cookei1211001000xanchus griseus1211001001torynchus cepedianus1011000001lamydoselachus anguineus1200001000

PAGE 255

Appendix E (continued): See Chapter 3 for abbreviations. 245 SpeciesCaCarcharhiCarcharhiCarcharhiNegaprion brevirostrisPrionace glaucaGaCentrophorusSqSquatina dumPrOxynotScymSomDaAculEtEtEchinorhinus cookeiHexanchus griseusNotChlam SoftMedHardSE LowerSE UpperLC LowerLC UpperLATMTrcharhinus perezi010121200nus leucas011221100nus plumbeus111121000nus limbatus010121100010022100110222100leocerdo cuvier011221100 granulosus110000100ualus acanthias110000000eril010002200istiophorus cirratum011001101us bruniensis???001110nodon ringens011002200niosus microcephalus010000100latias licha010021000eola nigra111001100mopterus virens100000110mopterus lucifer110000110110221110110220101orynchus cepedianus010200101ydoselachus anguineus100002220

PAGE 256

Appendix F: Continuous Variables for Evolutionary Analysis 246

PAGE 257

Appendix F: Heterodontus francisci.. See Chapter 3 for abbreviations. AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.481.212.002.502.3576.002.501.042.601.151.70172.30SD0.090.360.420.000.218.910.000.060.140.070.008.06Mean1.441.212.102.852.4569.552.551.102.801.151.65167.35SD0.270.410.420.070.3511.810.350.040.280.070.2111.53Mean1.411.131.952.352.2065.252.351.062.501.101.55162.35SD0.010.020.210.210.285.300.210.150.570.140.212.62Mean0.680.542.803.051.4578.601.601.622.601.500.90121.55SD0.320.160.711.200.074.380.140.120.420.710.0026.23Mean0.175.906.401.003.200.973.10SDMean0.175.405.500.902.900.972.80SDMean1.291.051.932.532.2592.002.501.002.501.201.55167.05SD0.180.000.380.570.075.370.000.060.140.000.218.84Mean1.411.191.932.372.2373.952.401.042.501.171.53166.03SD0.390.260.400.750.062.330.200.070.260.420.211.46Mean1.260.971.972.431.9079.302.071.102.271.131.40157.93SD0.300.180.120.250.2613.860.150.010.150.230.265.97Mean0.890.732.102.431.4096.401.601.442.151.250.90153.35SD0.680.470.400.470.570.570.550.070.640.2812.66Mean0.520.353.834.131.1792.152.071.382.631.650.85118.00SD0.020.181.891.710.387.001.000.380.640.210.078.77Mean0.210.235.475.731.172.701.433.331.900.40129.10SD0.052.372.120.251.570.530.81Lower, 5Lower, 7Lower, 8Upper, 5Upper, 1Upper, 7Upper, 8Upper, 2Upper, 4Lower, 1Lower, 2Lower, 4 247

PAGE 258

Appendix F (continued): Heterodontus francisci. BOMean9.26SD13.09Mean8.51SD12.04MeanSDMean16.08SD6.61Mean17.67SD17.10Mean14.60SD18.42Lower, 1-2Upper, 1-2Upper, 4-5Upper, 7-8Lower, 4-5Lower 7-8 248 AARCARCACINALower, 1-21.451.2072.801.05169.85Lower, 4-51.050.8071.951.35142.00Lower 7-80.200.001.00Upper, 1-21.351.1083.001.00166.55Upper, 4-51.100.8587.851.25155.65Upper, 7-80.350.3046.081.40123.55

PAGE 259

Appendix F (continued): Orectolobus maculatus. AARCARBCWBWCHCADCLCIMCLNCWNHNAMean2.351.687.529.349.930.009.601.1210.684.209.47153.80SD0.360.364.164.564.590.005.060.055.301.833.183.16Mean1.591.426.008.387.700.007.181.278.743.235.13134.80SD0.590.434.024.294.090.004.250.195.061.512.814.18Mean1.071.015.788.084.730.004.851.256.133.173.37119.97SD0.080.663.313.121.750.001.880.082.631.301.275.05Mean0.900.728.838.936.200.005.201.407.103.873.43118.73SD0.180.271.210.582.070.001.040.311.060.760.767.54Mean0.510.446.136.472.650.003.101.504.653.201.55111.65SD0.180.060.230.150.350.000.140.180.350.990.074.31Mean0.400.305.504.901.400.003.400.622.102.501.00117.50SD1.130.800.00Mean1.921.816.9810.0811.380.0011.301.0611.844.808.90142.50SD0.230.474.314.954.880.005.230.085.482.243.5615.37Mean1.411.195.468.125.720.005.501.237.063.364.58119.64SD0.200.414.134.593.220.003.100.234.661.521.874.76Mean1.120.944.857.805.270.004.731.276.072.872.87112.03SD0.420.783.774.781.560.002.760.143.911.621.584.10Mean1.280.814.857.533.530.003.781.305.002.733.30117.33SD0.220.363.913.622.120.002.340.083.321.671.7114.81Mean0.970.955.907.604.470.004.531.265.973.002.87113.83SD0.250.513.300.891.210.001.110.443.100.440.5714.35Mean0.960.884.255.283.350.003.731.214.452.472.30126.47SD0.090.291.971.330.860.001.260.191.411.501.2113.66Upper, 7Lower, 5Lower, 7Lower, 8Upper, 1Lower, 1Lower, 2Lower, 4Upper, 5Upper, 8Upper, 2Upper, 4 249

PAGE 260

Appendix F (continued): Orectolobus maculatus. BOMean-67.92SD14.62Mean2.99SD5.97Mean5.26SD9.12Mean-70.49SD41.78Mean0.53SD12.18Mean5.03SD10.07Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 250 AARCARCACINALower, 1-21.971.550.001.20144.30Lower, 4-50.990.860.001.32119.35Lower 7-80.460.370.001.06114.58Upper, 1-21.661.500.001.15131.07Upper, 4-51.200.870.001.29114.68Upper, 7-80.970.910.001.23120.15

PAGE 261

Appendix F (continued): Chiloscyllium plagiosum AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.140.9110.990.51110.70.8141.2SDMean1.251.330.911.2931.31.081.40.81162.3SDMean1.571.2111.21.40.931.30.71.1160SDMean1.3811.11.11.197.21.40.861.20.81.1143.1SDMean1.3310.90.90.9119.31.10.820.90.60.8149.2SDMean1.51.110.90.91961.20.8310.60.9163.1SDMean0.710.880.80.90.70.71.140.80.70.5160SDMean0.80.9SDMean1.170.811.10.80.81.381.10.60.7138.9SDMean10.9110.9106.511.21.20.80.8125.7SDMeanSDMeanSDLower, 2Lower, 4Upper, 5Upper, 8Upper, 2Upper, 4Upper, 7Lower, 5Lower, 7Lower, 8Upper, 1Lower, 1 251

PAGE 262

Appendix F (continued): Chiloscyllium plagiosum BOMean22.22SDMean9.52SDMeanSDMean10SDMean19.05SDMean22.22SDUpper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2Upper, 4-5 252 AARCARCINALower, 1-21.201.121.04151.75Lower, 4-51.471.100.89151.55Lower 7-81.421.060.83156.15Upper, 1-20.710.881.14160.00Upper, 4-51.080.851.29132.30Upper, 7-8

PAGE 263

Appendix F (continued): Stegostoma fasciatum AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.731.182.452.452.9049.353.200.993.151.302.25159.45SD0.160.080.070.070.280.640.420.020.350.000.212.33Mean2.552.60SD0.070.00Mean1.711.222.302.302.8064.953.001.023.051.252.15160.75SD0.190.040.280.280.424.450.570.080.350.070.350.49Mean1.421.212.602.553.1563.753.401.003.401.752.40158.40SD0.320.030.000.070.0711.100.000.000.000.490.148.34Mean1.251.102.102.152.3053.652.401.082.601.501.80153.05SD0.350.000.000.070.0010.820.000.000.000.420.000.78Mean1.291.212.152.252.6055.102.751.072.951.602.00161.20SD0.400.160.070.070.425.660.350.010.350.280.282.55Mean1.251.002.152.202.1052.102.301.002.301.201.50167.80SD0.070.00Mean1.340.932.202.202.0551.202.301.002.301.151.55146.95SD0.100.030.000.140.074.810.140.060.000.070.2125.95Mean1.210.912.402.452.2057.752.501.002.501.301.60159.45SD0.300.070.280.210.4218.880.280.060.420.140.5717.61Mean1.291.042.252.302.3060.602.551.002.551.251.60170.45SD0.190.160.350.420.0024.890.070.000.070.070.142.05Mean1.371.172.002.052.3565.402.501.002.501.201.65168.20SD0.130.160.140.210.4916.690.420.000.420.140.355.94Mean1.551.132.002.102.2552.152.451.022.501.101.70155.50SD0.260.110.000.000.210.070.350.030.280.000.287.64Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 253

PAGE 264

Appendix F (continued): Stegostoma fasciatum BOMean15.84SD0.22Mean14.15SD7.72Mean4.44SD6.29Mean9.10SD0.29Mean16.86SD0.75Mean14.30SD6.08Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 254 AARCARCACINALower, 1-21.731.1849.350.99159.45Lower, 4-51.571.2164.351.01159.58Lower 7-81.271.1554.381.08157.13Upper, 1-21.300.9751.651.00157.38Upper, 4-51.250.9759.181.00164.95Upper, 7-81.461.1558.781.01161.85

PAGE 265

Appendix F (continued): Ginglymostoma cirratum AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.000.535.706.033.0072.204.001.084.332.102.10140.97SD0.000.050.780.590.360.530.260.100.650.100.108.41Mean0.890.536.056.083.1783.604.171.064.402.402.10144.27SD0.150.020.400.400.3112.730.350.050.350.360.106.79Mean0.850.525.936.183.0582.884.051.124.502.632.15140.70SD0.170.051.021.040.5419.900.710.070.640.680.319.41Mean0.880.495.285.532.6375.773.551.124.002.201.93138.95SD0.160.070.920.910.696.130.700.090.930.590.499.31Mean0.960.525.335.532.7873.203.531.194.182.031.88137.15SD0.190.020.250.380.133.260.300.110.210.530.155.25Mean1.080.495.355.482.6068.183.301.284.201.651.78131.78SD0.080.020.650.690.353.290.480.140.390.170.132.40Mean0.870.494.785.142.3269.473.420.923.142.001.74144.30SD0.150.110.380.540.4515.410.400.080.340.200.336.11Mean0.790.485.145.522.4892.323.541.053.702.421.86135.62SD0.160.080.280.220.3716.170.400.130.270.580.2114.45Mean0.900.475.285.622.4273.623.421.133.862.001.80133.56SD0.120.090.981.020.3112.810.440.180.700.220.3212.60Mean0.920.455.806.042.6077.143.601.194.222.101.94128.66SD0.090.070.250.340.4210.570.470.140.190.120.268.89Mean0.820.415.425.842.2468.822.941.464.242.361.74132.46SD0.260.040.360.450.2936.980.380.210.421.040.2811.42Mean0.760.435.445.602.3074.353.001.404.132.281.63126.25SD0.190.060.740.960.489.660.620.230.651.030.4610.88Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 255

PAGE 266

Appendix F (continued): Ginglymostoma cirratum BOLower, 1-2Mean31.68SD12.80Lower, 4-5Mean15.66SD12.71Lower 7-8Mean15.66SD8.36Upper, 1-2Mean19.14SD4.03Upper, 4-5Mean20.05SD6.43Upper, 7-8Mean13.95SD4.54 256 AARCARCACINALower, 1-20.950.5377.901.07142.62Lower, 4-50.860.5079.321.12139.83Lower 7-81.020.5070.691.24134.46Upper, 1-20.830.4980.890.99139.96Upper, 4-50.910.4675.381.16131.11Upper, 7-80.790.4271.591.43129.36

PAGE 267

Appendix F (continued): Mitsukurina owstoni AARCARBCWBWCHCADCLCIMCLNCWNHNAMean5.093.516.89.923.9024.1124.23.517.8157.7SDMean5.692.728.712.723.7023.91.0525.13.922.2152.5SDMean3.948.117.203.212.6148SDMean3.881.998.415.716.70150.9714.53.312.8165.6SDMean6.814.30143.4SDMean3.211.897.414.513.20131.0613.83.310.6137.6SDMean5.182.847.411.52100.9720.93.417.6147.7SDMean4.122.238.811.819.6021.61.08203.918.4135.3SDMean3.441.687.916.213.3018.61.1516.43.211131.2SDMean7.515.9014.2137.7SDMean2.791.777.813.413.8014.11.1115.74.211.7134.4SDMean1.53915.913.8013.9113.9SDLower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 257

PAGE 268

Appendix F (continued): Mitsukurina owstoni BOMean0SDMean-25.55SDMean-17.75SDMean0SDMean-19.45SDMean-18.06SDUpper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 258 AARCARCACINALower, 1-25.393.120.001.03155.10Lower, 4-53.911.990.000.97156.80Lower 7-83.211.780.001.06140.50Upper, 1-24.952.530.001.02141.50Upper, 4-53.441.680.001.15134.45Upper, 7-82.791.650.001.06134.40

PAGE 269

Appendix F (continued): Alopias superciliosus AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.601.345.209.058.600.008.401.139.354.506.95140.35SD0.270.211.831.840.140.001.130.210.491.701.486.86Mean1.681.197.3010.239.330.008.801.1810.174.407.30148.60SD0.220.162.301.611.710.001.820.210.150.750.7015.21Mean1.241.067.4010.437.770.007.501.359.734.375.03125.77SD0.530.071.951.931.520.002.330.261.421.121.1011.58Mean1.030.976.9810.385.950.005.201.457.654.254.45127.85SD0.050.022.242.301.060.001.270.182.762.332.6211.81Mean2.000.866.979.806.200.004.701.366.402.204.40123.00SD0.252.330.00Mean0.920.805.438.974.350.003.801.726.354.053.55128.35SD0.360.100.152.030.350.000.570.681.631.060.493.75Mean1.991.656.8010.3010.100.006.901.8410.374.008.07143.80SD0.300.152.263.293.000.002.801.413.450.612.2915.00Mean1.811.447.2311.889.850.009.381.2011.434.257.35142.20SD0.500.382.732.832.460.001.810.183.491.251.3210.42Mean1.381.117.8311.938.630.007.981.4111.284.686.35123.58SD0.330.112.432.132.540.001.800.082.891.171.559.72Mean1.241.196.8011.738.300.008.131.3911.335.636.83120.27SD0.250.121.081.640.700.000.810.031.101.241.4510.99Mean1.020.856.3810.755.430.005.181.608.254.484.53337.03SD0.370.180.551.701.100.001.330.102.000.971.84433.60Mean0.840.865.659.855.170.005.171.638.374.273.53107.13SD0.190.311.242.251.270.001.270.191.821.100.9614.98Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 259

PAGE 270

Appendix F (continued): Alopias superciliosus BOMean-23.85SD3.11Mean-27.09SD9.72Mean-15.38SD3.67Mean-13.43SD10.86Mean-23.20SD4.60Mean-10.46SD8.08Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 260 AARCARCACINALower, 1-21.641.270.001.16144.48Lower, 4-51.141.020.001.40126.81Lower 7-81.460.830.001.54125.68Upper, 1-21.901.540.001.52143.00Upper, 4-51.311.150.001.40121.92Upper, 7-80.930.860.001.61222.08

PAGE 271

Appendix F (continued): Carcharias taurus AARCARBCWBWCHCADCLCIMCLNCWNHNAMean3.113.098.1015.4823.73136.8523.651.0123.956.5720.50162.33SD0.390.653.092.924.927.284.580.055.302.007.640.40Mean2.742.399.2516.1822.5797.8322.831.0323.606.8718.40146.47SD0.250.463.381.464.1359.634.460.035.352.314.565.51Mean2.772.707.3015.0714.0068.4013.651.0614.354.2511.85146.30SD0.080.673.662.174.1055.864.740.054.311.484.4511.17Mean2.311.936.8313.5310.2076.858.901.099.653.457.95141.00SD0.270.052.730.920.0044.341.130.030.920.070.787.64Mean1.781.954.478.708.607.501.158.603.205.70131.20SD1.300.70Mean1.511.134.857.505.4093.055.201.095.672.473.57139.37SD0.400.351.910.770.6160.170.850.191.440.570.2313.19Mean3.373.026.2811.1818.23114.6019.180.9217.754.3814.80169.73SD0.370.672.090.704.3343.784.100.064.641.646.297.82Mean3.122.678.7015.0819.3368.5318.631.0419.375.0716.00145.03SD0.620.253.281.695.4748.545.140.015.280.655.062.15Mean2.622.277.8516.7815.6776.9514.831.0615.674.6012.00145.25SD0.330.693.191.642.4153.532.150.012.040.420.421.34Mean2.271.967.4815.4512.90115.8012.751.1013.904.359.95130.20SD0.150.084.181.944.816.655.020.035.091.063.045.52Mean1.991.618.9815.8811.87113.2011.531.2113.805.0710.00125.20SD0.320.223.651.753.946.933.440.133.541.453.123.40Mean1.791.687.7014.2010.3072.809.571.2211.375.008.80133.40SD0.220.203.381.712.9549.363.070.142.381.561.702.12Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 261

PAGE 272

Appendix F (continued): Carcharias taurus BOMean1.46SD9.03Mean-31.43SD13.38Mean-16.50SD14.53Mean-5.19SD10.37MeanSDMean-17.73SD5.11Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 262 AARCARCACINALower, 1-22.922.74117.341.02154.40Lower, 4-52.542.3172.631.07143.65Lower 7-81.641.5493.051.12135.28Upper, 1-23.242.8591.570.98157.38Upper, 4-52.452.1196.381.08137.73Upper, 7-81.891.6593.001.21129.30

PAGE 273

Appendix F (continued): Pseudocarcharias kamoharai AARCARBCWBWCHCADCLCIMCLNCWNHNAMean4.344.183.555.3512.800.0012.071.0712.902.209.50166.00SD0.521.961.240.501.680.001.400.021.450.170.524.81Mean3.012.744.257.1510.830.0010.431.2112.632.807.87148.97SD0.881.171.350.610.760.000.290.111.230.950.062.49Mean2.102.302.705.935.000.004.701.055.002.004.20113.70SD0.700.970.510.820.000.440.231.45Mean1.941.212.985.334.270.004.131.325.431.703.27111.80SD0.230.121.120.400.380.000.450.170.570.200.156.68Mean1.260.763.554.002.700.002.251.573.452.803.3099.45SD0.260.090.210.140.140.000.490.380.071.701.412.05Mean1.802.600.00SDMean3.392.863.185.628.430.008.730.978.331.906.45138.80SD0.190.880.880.130.430.001.340.150.170.000.352.83Mean2.702.153.906.688.200.008.051.199.552.436.53138.90SD0.280.461.180.730.780.000.870.010.990.130.485.50Mean1.521.363.846.964.680.004.401.506.602.303.45100.45SD0.270.541.090.890.650.000.140.170.740.280.307.19Mean1.321.094.407.235.350.004.601.657.352.803.70100.75SD0.070.200.960.150.780.001.130.420.070.140.0013.51Mean0.820.883.636.373.170.002.471.974.832.872.2783.80SD0.160.160.351.590.310.000.150.290.460.780.319.01Mean0.710.514.105.172.070.002.172.455.002.401.6379.90SD0.150.110.891.590.550.000.830.941.470.980.495.82Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 263

PAGE 274

Appendix F (continued): Pseudocarcharias kamoharai BOMean-14.56SD14.21Mean-18.92SD17.51Mean0.00SD0.00Mean-9.49SD21.94Mean-17.53SD12.04Mean-15.54SD16.10Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 264 AARCARCACINALower, 1-23.683.460.001.14157.48Lower, 4-52.021.750.001.19112.75Lower 7-83.210.760.001.5799.45Upper, 1-23.052.500.001.08138.85Upper, 4-51.421.220.001.58100.60Upper, 7-80.760.700.002.2181.85

PAGE 275

Appendix F (continued): Isurus oxyrinchus AARCARBCWBWCHCADCLCIMCLNCWNHNAMean2.972.527.3610.9618.520.0018.341.0419.084.2412.30158.20SD0.290.271.302.153.550.003.270.042.891.352.4917.63Mean2.481.8211.3613.7019.660.0018.481.1220.826.2814.68154.02SD0.400.513.303.314.390.001.850.275.802.913.8413.56Mean1.290.9810.7413.7611.130.0011.201.1712.877.038.90144.60SD0.090.111.451.662.350.002.650.141.972.822.937.83Mean1.370.9710.4813.0410.030.0010.181.1511.735.307.18136.28SD0.120.071.641.492.160.002.050.052.411.341.5414.40Mean1.080.928.409.528.300.008.971.059.475.405.67138.40SD0.330.051.501.480.960.001.010.081.501.181.602.18Mean1.231.006.767.766.740.006.781.107.463.304.02136.30SD0.080.081.581.491.580.001.290.121.620.940.977.04Mean2.431.7610.1813.7418.040.0017.741.1720.645.6013.62143.02SD0.230.121.331.633.450.003.710.074.250.682.396.69Mean1.711.4611.3215.7216.500.0016.321.3121.327.6812.76139.24SD0.220.062.002.303.160.003.350.063.862.502.376.84Mean1.061.069.5415.149.600.009.271.5314.177.007.33131.23SD0.070.112.092.481.710.002.420.083.931.591.145.27Mean1.201.0510.7814.6612.530.0011.301.4716.707.779.29134.83SD0.040.121.662.062.460.001.840.113.481.811.838.48Mean1.231.0210.0812.5810.430.009.831.2812.535.706.88140.10SD0.150.081.882.411.820.001.930.042.471.401.083.68Mean1.050.908.5010.787.130.007.131.268.984.184.33130.53SD0.120.091.501.860.300.000.190.040.381.070.981.39Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 265

PAGE 276

Appendix F (continued): Isurus oxyrinchus BOMean-2.54SD5.67Mean-30.74SD12.94Mean-8.30SD11.38Mean-2.40SD5.37Mean-31.40SD12.34Mean-10.45SD6.42Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 266 AARCARCACINALower, 1-22.732.170.001.08156.11Lower, 4-51.330.980.001.16140.44Lower 7-81.150.960.001.08137.35Upper, 1-22.071.610.001.24141.13Upper, 4-51.131.050.001.50133.03Upper, 7-81.140.960.001.27135.31

PAGE 277

Appendix F (continued): Carcharodon carcharias AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.991.6012.6618.7820.050.0020.950.9720.387.4314.55162.10SD0.180.071.743.372.530.002.570.062.681.642.214.94Mean1.481.3914.9820.5022.950.0023.301.0223.7010.4015.35159.75SD0.010.214.273.205.440.005.800.035.232.693.893.18Mean1.391.1213.5418.3214.500.0016.331.0016.187.1810.18160.08SD0.590.193.903.942.140.003.630.072.452.065.549.78Mean1.481.1513.2617.5414.920.0015.921.0015.927.2210.34159.18SD0.190.143.282.671.600.002.340.041.822.181.692.62Mean1.291.1310.1212.489.700.009.271.2011.104.706.07150.63SD0.100.152.532.030.400.000.960.050.891.301.657.25Mean1.340.938.989.826.900.007.471.057.834.105.17157.47SD0.240.222.582.490.360.000.060.090.682.001.4423.67Mean1.421.3616.4319.1022.200.0022.701.0323.3314.1019.90169.53SD0.070.073.332.863.430.003.370.043.023.013.478.69Mean1.241.1819.5621.6624.900.0026.401.0527.7012.8018.35176.40SD0.080.195.494.795.030.006.350.035.953.901.205.09Mean1.071.0219.3224.5619.400.0018.951.2323.5013.2814.10154.95SD0.130.124.845.243.500.003.510.085.813.413.176.08Mean1.031.0020.6023.2220.400.0021.161.1524.1013.9014.24160.40SD0.130.075.584.584.390.004.410.074.033.363.382.43Mean1.080.9416.9819.2815.860.0016.921.1218.2610.7011.40154.10SD0.080.064.533.963.640.005.100.234.233.663.093.97Mean1.040.8713.9415.4011.750.0011.531.2514.357.657.85142.85SD0.130.063.243.932.460.002.600.063.182.141.644.57Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 267

PAGE 278

Appendix F (continued): Carcharodon carcharias BOMean-21.62SD5.81Mean-30.88SD10.88Mean-15.94SD5.50Mean-4.17SD8.33Mean-5.96SD5.46Mean-11.72SD3.98Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 268 AARCARCACINALower, 1-21.731.500.001.00160.93Lower, 4-51.431.140.001.00159.63Lower 7-81.311.030.001.12154.05Upper, 1-21.331.270.001.04172.96Upper, 4-51.051.010.001.19157.68Upper, 7-81.060.900.001.18148.48

PAGE 279

Appendix F (continued): Lamna nasus AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.561.606.679.839.7595.9010.250.979.854.807.40150.95SD0.530.432.603.153.0432.104.170.063.460.282.121.34Mean2.112.156.7710.637.307.701.048.001.803.80142.60SD3.623.23Mean1.491.276.179.537.2097.958.251.027.954.306.35139.10SD0.060.323.513.584.2429.634.880.203.321.842.478.34Mean1.631.526.609.707.2576.408.200.937.754.307.00132.00SD0.926.084.113.181.840.163.04Mean1.641.187.0010.307.0582.357.151.077.603.956.40135.25SD0.350.593.960.570.497.711.340.100.710.490.576.43Mean1.221.396.309.607.1574.056.651.036.904.205.20140.05SD0.350.734.382.121.483.461.770.021.980.421.9814.21Mean1.942.034.939.778.75129.609.301.019.404.158.15142.50SD0.140.321.953.293.752.970.002.971.202.905.80Mean1.621.625.4310.336.00118.706.401.117.103.706.00180.00SD1.752.98Mean1.551.324.5310.476.1037.406.201.086.853.805.90114.00SD0.420.763.993.252.260.123.18Mean1.281.205.4710.534.804.401.305.702.503.20149.80SD1.313.89Mean1.421.295.478.436.6585.306.101.116.903.304.80142.60SD0.220.161.363.762.9017.252.830.093.680.992.124.53Mean1.741.216.2011.057.4561.807.051.158.103.205.55132.05SD0.110.080.571.340.210.490.040.850.570.644.88Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 269

PAGE 280

Appendix F (continued): Lamna nasus BOMean-21.70SD32.98Mean-20.02SD38.25Mean5.65SD20.56Mean-13.29SD33.09Mean-18.39SD29.54Mean-4.19SD5.92Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 270 AARCARCACINALower, 1-21.831.8795.901.01146.78Lower, 4-51.561.4087.180.98135.55Lower 7-81.431.2878.201.05137.65Upper, 1-21.781.83124.151.06161.25Upper, 4-51.421.2637.401.19131.90Upper, 7-81.581.2573.551.13137.33

PAGE 281

Appendix F (continued): Scyliorhinus retifer AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.371.070.850.950.9070.001.001.051.050.550.75133.75SD0.050.100.210.210.140.140.060.210.070.073.18Mean1.701.041.031.171.0757.201.231.021.230.570.93142.87SD0.780.310.060.210.290.320.160.250.060.324.06Mean1.330.851.001.180.8379.631.031.121.150.550.73150.48SD0.200.250.220.210.2212.460.170.050.170.130.157.89Mean1.570.941.031.130.9757.151.101.081.200.570.83134.83SD0.590.210.150.230.2524.110.200.080.300.210.219.73MeanSDMeanSDMean1.431.001.051.131.0867.631.181.001.200.600.88146.55SD0.200.180.190.170.3626.120.360.110.420.180.3313.42Mean1.471.100.871.020.9783.801.001.071.070.530.80127.97SD0.320.170.150.130.310.990.300.070.310.120.3014.07Mean1.240.901.101.270.9759.051.001.371.300.670.83112.17SD0.100.270.300.420.3111.100.360.340.260.250.3515.24Mean1.500.890.901.400.800.801.381.100.400.60133.30SDMean1.580.730.850.900.6066.800.801.220.950.450.70134.45SD0.120.180.210.000.280.160.210.210.2816.76Mean1.400.621.301.500.8069.500.801.501.200.500.70103.80SDLower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 271

PAGE 282

Appendix F (continued): Scyliorhinus retifer BOMeanSDMeanSDMeanSDMeanSDMeanSDMeanSDUpper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 272 AARCARCACINALower, 1-21.531.0663.601.03138.31Lower, 4-51.450.8968.391.10142.65Lower 7-8Upper, 1-21.451.0575.721.04137.26Upper, 4-51.370.8959.051.37122.73Upper, 7-81.490.6768.151.36119.13

PAGE 283

Appendix F (continued): Galeus arae AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.501.250.400.500.5042.300.501.000.500.200.30160.00SDMean2.502.000.300.400.6075.300.601.000.600.200.50177.90SDMean2.001.670.300.400.50107.400.601.000.600.200.40158.10SDMean2.001.250.400.500.5058.300.501.200.600.200.4067.10SDMeanSD138.10Mean2.001.250.400.400.5064.000.501.000.500.200.40SD150.20Mean1.501.000.500.500.5069.300.501.000.500.200.30SD143.10Mean1.671.200.500.600.6060.400.601.000.600.300.50SDMean1.331.200.500.700.6057.900.700.860.600.300.40164.80SDMeanSDMeanSDMeanSDLower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 273

PAGE 284

Appendix F (continued): Galeus arae BOMeanSDMean-22.22SDMean-50.00SDMean-18.18SDMeanSDMeanSDUpper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 274 AARCARCACINALower, 1-22.001.6358.801.00168.95Lower, 4-52.001.4682.851.10112.60Lower 7-82.001.2564.001.00138.10Upper, 1-21.581.1064.851.00146.65Upper, 4-51.331.2057.900.86164.80Upper, 7-8

PAGE 285

Appendix F (continued): Paramaturus xaniurus AARCARBCWBWCHCADCLCIMCLNCWNHNAMeanSDMeanSDMeanSDMeanSDMeanSDMeanSDMean1.000.630.800.900.5064.000.601.170.700.500.50135.90SDMean1.000.710.701.000.5067.300.601.000.600.400.40142.90SDMean1.000.710.700.800.5063.600.601.000.600.400.40136.70SDMeanSDMeanSDMeanSDLower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 275

PAGE 286

Appendix F (continued): Paramaturus xaniurus BOMeanSDMeanSDMeanSDMeanSDMeanSDMeanSDUpper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 276 AARCARCACINALower, 1-2Lower, 4-5Lower 7-8Upper, 1-21.000.6365.651.08139.40Upper, 4-51.000.7163.601.00136.70Upper, 7-8

PAGE 287

Appendix F (continued): Pseudotriakis microdon AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.131.130.800.900.900.001.001.001.000.800.90180.00SDMean1.331.330.600.600.800.000.901.000.900.600.80180.00SDMean1.501.500.800.801.200.001.301.001.300.801.20180.00SDMean1.861.860.700.801.300.001.301.081.400.701.30180.00SDMean1.001.001.001.201.000.001.101.091.201.001.00180.00SDMeanSDMean1.501.220.900.901.1093.601.300.921.200.600.90160.80SDMean1.001.00SDMean1.711.501.001.301.500.001.700.881.500.701.20166.20SDMean1.671.500.801.001.200.001.300.921.200.601.00173.00SDMean1.501.220.901.101.100.001.300.921.200.600.90170.50SDMean1.201.30SDLower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 277

PAGE 288

Appendix F (continued): Pseudotriakis microdon BOMean-10.53SDMean0.00SDMean0.00SDMean-13.33SDMean0.00SDMean0.00SDUpper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 278 AARCARCACINALower, 1-21.231.230.001.00180.00Lower, 4-51.681.680.001.04180.00Lower 7-81.001.000.001.09180.00Upper, 1-21.501.0046.800.92160.80Upper, 4-51.691.500.000.90169.60Upper, 7-81.501.220.000.92170.50

PAGE 289

Appendix F (continued): Triakis semifasciata AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.000.862.903.202.5091.302.601.233.202.102.10126.80SDMean1.280.793.403.702.70101.703.001.173.501.802.30116.30SDMean1.200.693.503.802.402.401.633.901.501.80119.40SDMean1.310.773.103.802.402.401.503.601.301.70138.00SDMean0.930.583.804.402.202.101.904.001.501.40120.90SDMean1.200.553.804.002.1058.802.201.773.901.001.20104.70SDMean1.300.952.002.001.90111.002.200.952.101.001.30150.10SDMean1.440.952.002.401.902.200.912.000.901.30159.00SDMean1.621.252.002.202.5070.602.601.042.701.302.10158.70SDMean1.430.922.502.502.302.800.932.601.402.00137.40SDMean1.200.852.602.802.202.401.132.701.501.80138.70SDMean1.000.733.003.102.2092.702.301.353.101.701.70116.20SDLower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 279

PAGE 290

Appendix F (continued): Triakis semifasciata BOMean23.19SDMean18.42SDMean19.05SDMean31.82SDMean17.02SDMean23.73SDUpper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 280 AARCARCACINALower, 1-21.140.8396.501.20121.55Lower, 4-51.250.731.56128.70Lower 7-81.070.5758.801.84112.80Upper, 1-21.370.95111.000.93154.55Upper, 4-51.521.0970.600.98148.05Upper, 7-81.100.7992.701.24127.45

PAGE 291

Appendix F (continued): Carcharhinus leucas AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.691.1010.1411.4211.040.0011.281.0812.305.349.04135.46SD0.220.082.372.221.920.002.250.073.010.901.936.95Mean1.401.0812.6614.5813.900.0015.201.0816.438.4311.03144.93SD0.330.021.962.212.930.002.720.103.043.521.472.90Mean1.470.9515.4416.6215.000.0016.151.0817.337.4510.80138.65SD0.170.082.932.911.780.002.860.101.591.581.039.97Mean1.360.9115.4816.8613.970.0014.931.1316.908.0010.80137.70SD0.160.103.503.292.290.003.010.023.581.351.134.97Mean1.340.8015.5816.9012.440.0014.041.1015.386.768.92134.14SD0.160.071.871.971.210.001.720.031.601.340.927.75Mean1.180.7114.9216.3011.600.0013.551.1515.756.657.80135.15SD0.170.012.362.222.120.002.050.124.030.210.850.64Mean1.151.0612.8413.4813.850.0014.851.0615.638.659.95162.65SD0.060.162.382.271.640.002.420.111.831.642.015.31Mean1.051.0316.0017.7416.240.0017.701.0518.609.9410.36167.68SD0.050.113.753.721.940.002.600.102.821.761.361.90Mean1.020.8020.5622.1016.340.0017.921.2321.8411.2511.30152.65SD0.110.043.004.001.700.002.860.102.952.501.832.82Mean0.950.7720.6421.7215.900.0017.141.2821.8810.9410.38150.92SD0.040.052.412.441.330.002.000.061.990.710.882.82Mean0.890.7420.2821.5615.000.0016.221.2620.3610.649.40146.78SD0.070.031.292.111.270.001.050.101.631.330.574.42Mean0.850.7220.1822.1414.440.0015.021.3420.2211.359.68146.23SD0.050.062.572.801.310.001.490.052.450.570.546.72Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 281

PAGE 292

Appendix F (continued): Carcharhinus leucas BOMean6.38SD9.04Mean6.15SD6.33Mean6.46SD7.01Mean22.52SD3.16Mean22.04SD12.65Mean21.58SD3.58Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 282 AARCARCACINALower, 1-21.551.090.001.08140.20Lower, 4-51.420.930.001.11138.18Lower 7-81.260.750.001.13134.65Upper, 1-21.101.050.001.06165.17Upper, 4-50.980.790.001.25151.79Upper, 7-80.870.730.001.30146.50

PAGE 293

Appendix F (continued): Carcharhinus limbatus AARCARBCWBWCHCADCLCIMCLNCWNHNAMean2.471.198.159.359.300.0010.200.919.303.007.40120.40SD0.490.780.00Mean2.001.029.7011.1010.200.0010.501.0511.003.858.60123.70SD0.420.420.000.64Mean1.730.959.8510.209.350.0010.800.9810.554.607.95131.30SD0.070.040.351.980.070.000.140.010.210.140.073.25Mean1.660.8610.3011.508.750.0010.501.0711.304.807.70127.40SD0.360.140.991.270.640.000.140.232.551.560.852.55Mean1.410.7510.3011.558.300.0010.601.2413.104.658.30131.30SD0.991.200.001.77Mean1.840.799.7510.157.650.009.100.999.003.756.90129.25SD0.020.040.210.920.210.000.140.020.280.350.571.63Mean1.551.176.807.307.800.008.650.907.654.206.50140.70SD0.070.221.270.570.000.000.920.200.920.000.285.37Mean1.481.098.0010.208.700.008.801.039.105.207.70138.50SD0.00Mean1.570.8111.6512.9510.600.0011.301.0511.904.607.70129.30SD2.050.920.00Mean1.310.8212.6013.7510.250.0010.351.2012.406.258.00125.70SD0.190.071.272.190.210.000.070.010.001.771.130.14Mean1.540.7312.7013.659.200.009.851.2512.304.556.70125.35SD0.410.061.702.190.420.000.350.070.281.480.420.64Mean1.220.7212.4013.808.900.00938.000.0111.906.007.30117.20SDUpper, 8Upper, 2Upper, 4Upper, 5Upper, 7Lower, 5Lower, 7Lower, 8Upper, 1Lower, 1Lower, 2Lower, 4 283

PAGE 294

Appendix F (continued): Carcharhinus limbatus BOMean1.20SD1.70Mean3.25SD0.07Mean1.65SD0.35Mean2.90SDMean3.60SD0.57Mean4.00SDUpper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 284 AARCARCACINALower, 1-22.231.110.000.98122.05Lower, 4-51.700.900.001.03129.35Lower 7-81.620.770.001.11130.28Upper, 1-21.511.130.000.96139.60Upper, 4-51.440.810.001.13127.50Upper, 7-81.380.720.000.63121.28

PAGE 295

Appendix F (continued): Carcharhinus perezi AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.340.908.139.706.930.007.500.997.434.005.35125.65SD0.060.010.730.750.590.000.980.081.230.280.640.92Mean1.690.989.5211.529.150.009.881.0410.154.437.48130.03SD0.100.070.850.740.540.001.290.090.560.210.534.94Mean1.620.8611.4212.639.820.0011.071.0611.654.737.68131.68SD0.080.070.980.800.640.001.000.100.740.280.558.66Mean1.550.8312.6013.0010.080.0011.081.0511.585.027.70132.58SD0.180.071.331.890.990.001.000.040.890.650.777.41Mean1.420.7611.3813.228.550.009.751.0510.254.606.53122.33SD0.080.151.531.590.440.000.700.070.830.320.594.33Mean1.300.6911.5813.258.070.009.330.999.234.636.03121.60SD0.080.041.301.611.240.001.210.080.760.710.783.57Mean1.441.038.3010.178.480.009.151.019.284.376.28369.80SD0.090.100.941.390.770.000.560.040.830.390.31535.90Mean1.611.0710.0211.9710.580.0010.421.1011.404.787.36142.54SD0.420.120.731.071.320.000.870.101.221.060.643.86Mean1.120.8014.2315.7011.430.0011.171.2714.126.787.58135.86SD0.070.041.661.771.380.001.270.091.290.980.795.92Mean1.091.1113.2016.0310.000.0010.331.3013.376.376.93133.80SD0.060.624.072.030.300.001.110.110.320.250.312.85Mean0.980.8713.4815.3211.380.009.621.4313.726.766.60124.20SD0.060.372.392.153.860.001.130.051.500.940.983.97Mean0.880.6812.7814.588.570.008.631.5713.436.906.00121.43SD0.120.111.651.470.210.000.850.201.171.130.269.27Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 285

PAGE 296

Appendix F (continued): Carcharhinus perezi BOMean16.08SD9.69Mean13.09SD4.42Mean14.74SD5.17Mean21.14SD5.83Mean16.96SD3.26Mean27.33SD3.70Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 286 AARCARCACINALower, 1-21.510.940.001.01127.84Lower, 4-51.590.850.001.05132.13Lower 7-81.360.730.001.02121.96Upper, 1-21.531.050.001.05256.17Upper, 4-51.110.960.001.29134.83Upper, 7-80.930.770.001.50122.82

PAGE 297

Appendix F (continued): Carcharhinus plumbeus AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.770.976.977.436.670.007.031.047.303.475.97135.77SD0.430.131.460.380.760.000.740.051.050.670.653.44Mean1.750.938.4310.508.050.008.800.998.604.006.90131.05SD0.310.080.681.440.070.001.270.210.570.710.005.73Mean1.560.869.7011.138.550.009.800.959.304.006.25134.45SD0.020.010.690.500.640.000.710.030.420.000.075.16Mean1.330.8610.2010.638.700.009.431.029.605.337.10127.27SD0.140.140.791.400.720.000.230.121.280.380.879.92Mean1.220.6610.2711.476.730.007.871.048.174.435.37127.10SD0.050.040.900.810.210.000.210.100.640.680.575.37Mean1.440.6410.3310.736.600.007.971.098.673.575.07128.10SD0.290.030.380.670.170.000.490.130.470.380.571.51Mean1.291.107.908.708.600.009.031.039.335.306.83156.57SD0.050.151.110.790.260.000.210.040.400.200.126.67Mean1.130.9311.0712.5010.070.0010.501.1411.977.278.23148.57SD0.050.162.061.470.850.000.530.030.570.210.598.72Mean1.000.7713.0314.0710.000.0010.731.3013.737.207.23137.87SD0.040.050.810.810.000.002.070.200.810.440.657.46Mean0.930.7413.4015.439.800.0010.031.4113.908.007.47133.53SD0.110.091.711.700.100.001.550.220.460.661.0611.82Mean0.830.7312.3713.978.850.008.601.5213.006.555.45139.25SD0.010.050.760.810.070.000.570.281.561.341.203.46Mean0.750.6912.4513.158.500.008.851.4712.858.256.15124.20SD0.050.141.341.630.850.000.920.351.770.490.0711.17Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 287

PAGE 298

Appendix F (continued): Carcharhinus plumbeus BOMean18.07SD11.83Mean15.62SD15.98Mean13.69SD12.25Mean24.68SD9.94Mean17.88SD8.68Mean31.01SD2.65Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 288 AARCARCACINALower, 1-21.760.950.001.01133.41Lower, 4-51.450.860.000.98130.86Lower 7-81.330.650.001.07127.60Upper, 1-21.211.010.001.09152.57Upper, 4-50.970.750.001.36135.70Upper, 7-80.790.710.001.50131.73

PAGE 299

Appendix F (continued): Negaprion brevirostris AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.941.409.7828.8713.940.0013.241.1515.185.9011.33421.85SD0.170.141.8040.161.940.001.420.111.611.141.49604.85Mean1.781.0611.7513.8312.240.0013.621.1215.266.2210.98127.98SD0.090.241.732.152.550.001.910.062.221.081.434.65Mean1.670.9112.5314.3311.340.0012.161.0512.705.409.00120.86SD0.060.122.292.251.700.001.990.052.131.101.674.09Mean1.530.8812.5714.2812.100.0012.471.1614.406.309.53118.07SD0.180.081.872.160.100.001.350.021.570.950.750.70Mean1.410.7312.5413.849.470.0010.501.0811.534.736.60115.70SD0.090.102.983.042.020.002.420.113.721.752.264.45Mean1.350.6911.5813.748.000.008.331.2610.584.406.00118.05SD0.100.042.532.531.930.001.620.152.961.031.665.25Mean1.801.338.4710.1511.170.0011.551.0412.035.339.50131.13SD0.210.141.681.711.800.001.900.042.360.981.529.20Mean1.761.299.4710.9711.980.0011.281.1613.155.8310.37136.20SD0.160.111.071.502.530.002.050.032.600.992.4819.24Mean1.220.8116.5417.7613.900.0014.571.2418.038.9310.97118.50SD0.060.032.682.472.250.002.740.023.232.353.334.16Mean1.310.7616.3717.4812.620.0012.441.4117.506.588.63120.33SD0.140.052.612.692.050.001.950.062.411.181.845.13Mean1.200.7814.1815.4311.220.0011.081.4616.226.657.95116.10SD0.070.082.443.162.340.001.380.072.791.792.042.08Mean1.100.7713.3315.0310.220.0010.061.3512.966.737.38112.03SD0.100.061.922.641.630.002.240.443.931.421.484.17Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 289

PAGE 300

Appendix F (continued): Negaprion brevirostris BOMean9.71SD9.39Mean7.96SD5.41Mean9.73SD6.71Mean12.99SD11.45Mean14.77SD4.79Mean15.11SD8.91Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 290 AARCARCACINALower, 1-21.861.230.001.14274.92Lower, 4-51.600.890.001.10119.46Lower 7-81.380.710.001.17116.88Upper, 1-21.781.310.001.10133.67Upper, 4-51.260.780.001.33119.41Upper, 7-81.150.780.001.40114.06

PAGE 301

Appendix F (continued): Galeocerdo cuvier AARCARBCWBWCHCADCLCIMCLNCWNHNAMean0.710.7416.8518.6012.6088.2513.801.2215.9010.157.13127.75SD0.090.103.484.213.5614.004.480.344.082.731.6615.24Mean0.700.7018.9019.1813.4674.2415.581.2316.8210.647.32119.22SD0.060.084.975.724.4912.197.580.464.943.482.0515.40Mean0.630.6621.2821.5613.2874.8014.801.3819.4810.606.60112.00SD0.050.056.286.134.8912.876.550.215.903.701.9119.31Mean0.500.6021.9422.8213.5074.5515.171.4521.6012.335.73104.80SD0.110.017.007.834.2017.895.300.206.785.251.5016.26Mean0.460.5419.6020.5611.6077.2812.251.6519.9011.585.33105.03SD0.050.036.287.142.996.873.670.105.183.191.5214.45Mean0.420.4918.5018.369.9569.6311.881.4817.0011.374.73109.55SD0.050.026.095.922.6412.564.140.184.491.030.238.48Mean0.730.7420.0620.5816.4885.0317.701.3123.3512.058.85127.15SD0.050.057.187.134.797.974.890.096.733.532.8813.21Mean0.670.7221.4022.1215.5672.8217.501.2319.8811.347.56112.84SD0.070.066.347.335.529.847.710.326.133.962.4910.63Mean0.660.7022.3223.3615.3873.1415.921.4222.1211.647.52116.44SD0.080.067.476.984.8817.175.360.186.223.882.1414.37Mean0.600.6322.8224.5815.6076.4316.431.4523.6812.857.48114.20SD0.070.096.917.835.027.745.160.136.743.971.859.43Mean0.490.5519.4619.5211.6065.1312.181.6319.6512.155.90100.63SD0.040.026.276.112.9712.593.280.124.883.571.5911.92Mean0.410.4916.9617.848.3662.428.801.8115.829.663.8892.42SD0.060.054.865.992.857.782.720.074.533.291.007.91Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 291

PAGE 302

Appendix F (continued): Galeocerdo cuvier BOMean8.91SD6.12Mean2.67SD3.82Mean-1.05SD4.26Mean6.06SD6.34Mean3.54SD8.04Mean1.97SD12.69Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 292 AARCARCACINALower, 1-20.710.7281.251.23123.49Lower, 4-50.560.6374.681.41108.40Lower 7-80.440.5273.451.56107.29Upper, 1-20.700.7378.921.27120.00Upper, 4-50.630.6674.781.44115.32Upper, 7-80.450.5263.771.7296.52

PAGE 303

Appendix F (continued): Prionace glauca AARCARBCWBWCHCADCLCIMCLNCWNHNAMean2.265.186.638.9732.080.0012.421.0312.904.6310.33148.18SD0.317.651.471.5444.290.002.120.112.901.052.199.22Mean7.241.757.879.2214.300.0013.761.1115.324.2832.08143.76SD10.200.171.611.631.710.001.730.052.000.6947.486.54Mean2.081.518.9010.3012.230.0011.651.1613.484.759.60133.40SD0.350.061.451.091.180.001.210.222.731.291.438.09Mean1.651.1910.0511.0811.350.0011.201.1312.485.088.18129.28SD0.310.111.461.611.230.001.330.272.211.081.166.86Mean1.460.9210.4211.9310.230.009.981.2612.554.887.15121.98SD0.170.062.042.661.600.001.430.091.660.671.261.41Mean1.610.909.7011.129.500.0010.031.1411.454.407.05126.85SD0.180.092.212.171.010.000.280.232.430.631.124.08Mean1.421.1312.2413.3814.130.0014.851.1216.237.4810.53138.63SD0.200.090.811.350.920.001.920.271.810.850.558.12Mean1.161.0112.6713.9312.600.0012.131.4117.177.438.60135.00SD0.040.152.192.151.420.001.210.082.121.001.1511.98Mean1.120.9512.9414.2211.750.0011.951.4416.606.937.75128.08SD0.190.081.971.912.310.003.110.251.770.741.424.14Mean0.970.8812.7514.5510.660.0011.821.3115.247.827.58334.68SD0.170.112.312.582.120.002.040.374.101.111.50470.89Mean0.930.8212.2813.969.980.0011.541.2213.466.986.46116.68SD0.090.052.132.691.270.001.730.534.760.981.073.75Mean0.890.9412.1514.1310.600.0010.421.3413.426.926.10118.30SD0.110.262.292.590.680.001.630.423.190.790.423.45Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 293

PAGE 304

Appendix F (continued): Prionace glauca BOMean8.93SD9.96Mean6.91SD12.24Mean3.38SD14.77Mean27.78SD1.37Mean21.57SD6.50Mean23.49SD3.83Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 294 AARCARCACINALower, 1-24.753.470.001.07145.97Lower, 4-51.861.350.001.15131.34Lower 7-81.540.910.001.20124.41Upper, 1-21.291.070.001.27136.81Upper, 4-51.050.910.001.37231.38Upper, 7-80.910.880.001.28117.49

PAGE 305

Appendix F (continued): Sphyrna mokarran AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.421.0810.0311.9510.720.0011.061.0911.846.108.64140.52SD0.090.091.421.701.010.001.960.151.050.530.8514.12Mean1.401.1411.4312.5713.080.0013.281.0113.436.859.63145.50SD0.030.101.491.530.760.001.410.091.370.410.7115.86Mean1.311.0511.9513.3012.980.0012.751.3116.537.7510.18137.60SD0.020.091.381.471.590.001.810.111.480.801.1313.50Mean1.180.9511.8213.2810.580.0011.501.1012.456.988.25134.00SD0.110.071.471.331.580.001.650.242.440.741.547.63Mean1.140.9211.6812.7810.480.0010.301.2612.906.627.58137.16SD0.120.061.241.661.500.001.630.101.690.701.227.91Mean1.140.8711.7212.7010.180.0010.671.2413.136.637.55133.47SD0.170.081.031.161.570.002.080.101.920.551.3313.30Mean1.140.928.078.936.800.007.231.117.854.334.93139.03SD0.070.231.531.281.340.001.360.190.970.700.7410.58Mean0.930.8210.2010.457.680.007.351.5111.006.085.68117.20SD0.050.182.131.901.630.001.420.141.850.831.0312.92Mean0.950.7911.8212.758.780.008.881.5513.786.756.45117.88SD0.110.142.812.331.610.001.160.132.101.331.505.10Mean0.880.8012.2212.729.480.008.581.7314.507.506.60127.55SD0.130.071.211.061.470.002.080.251.650.481.2512.12Mean0.940.7313.2714.1310.030.009.651.6616.038.087.58114.73SD0.060.091.211.271.240.000.790.071.120.590.192.67Mean0.880.6713.1013.629.100.0039.731.1715.207.937.00117.30SD0.090.091.080.871.560.0053.930.881.500.350.406.03Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 295

PAGE 306

Appendix F (continued): Sphyrna mokarran 296UULLU BOMean20.54SD40.43Mean21.61SD67.12Mean22.24SD45.40Mean25.03SD16.14Mean15.67SD70.59Mean12.69SD117.84pper, 4-5pper, 7-8ower, 1-2ower, 4-5Lower 7-8pper, 1-2 AARCARCACINALower, 1-21.411.110.001.05143.01Lower, 4-51.241.000.001.20135.80Lower 7-81.140.890.001.25135.31Upper, 1-21.040.870.001.31128.11Upper, 4-50.920.790.001.64122.71Upper, 7-80.910.700.001.42116.01

PAGE 307

Appendix F (continued): Sphyrna tiburo AARCARBCWBWCHCADCLCIMCLNCWNHNAMean0.940.563.844.162.12111.002.641.173.101.641.52132.74SD0.160.070.620.620.150.110.180.520.270.2211.92Mean0.750.504.704.602.362.861.403.862.521.82123.16SD0.160.040.220.530.150.520.370.360.700.237.68Mean0.540.325.225.841.6485.702.321.854.242.381.28112.84SD0.180.060.640.960.324.190.400.160.430.190.488.79Mean0.460.355.245.601.8295.142.381.744.142.161.00122.28SD0.070.040.460.620.2714.500.250.110.360.110.1411.45Mean0.300.285.185.561.48106.902.161.843.901.880.58131.38SD0.120.080.360.630.4613.300.390.210.310.130.266.74Mean0.220.244.845.381.14126.702.021.753.481.900.40140.85SD0.050.060.190.300.2332.770.240.330.290.450.0819.75Mean0.690.512.983.521.5280.901.701.572.681.400.96116.10SD0.150.050.380.520.190.000.140.200.440.290.2111.07Mean0.550.423.924.381.7385.751.901.993.751.951.05102.68SD0.090.060.770.880.4614.640.420.160.720.650.3111.59Mean0.580.454.845.282.1465.432.321.954.522.201.20104.48SD0.160.060.901.060.2120.930.220.300.770.690.147.71Mean0.490.356.227.042.2579.802.582.215.682.731.3394.38SD0.030.010.990.910.4010.800.450.050.930.560.267.12Mean0.370.355.726.241.9661.772.262.245.022.781.0294.72SD0.080.050.680.690.1917.820.250.250.420.340.1810.32Mean0.320.345.406.101.8466.652.002.464.923.120.9491.02SD0.120.050.770.660.1710.560.160.360.780.610.177.16Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 297

PAGE 308

Appendix F (continued): Sphyrna tiburo BOMean23.55SD9.39Mean22.07SD3.64Mean23.35SD4.11Mean28.47SD5.05Mean24.07SD7.45Mean24.45SD7.80Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 298 AARCARCACINALower, 1-20.850.530.001.29127.95Lower, 4-50.500.3390.421.80117.56Lower 7-80.260.26116.801.79136.12Upper, 1-20.620.4783.331.78109.39Upper, 4-50.540.4072.622.0899.43Upper, 7-80.340.3564.212.3592.87

PAGE 309

Appendix F (continued): Hemipristis elongata AARCARBCWBWCHCADCLCIMCLNCWNHNAMean4.684.651.742.448.0763.707.670.997.631.707.67174.37SD0.971.020.610.961.080.490.050.860.460.409.58Mean3.543.032.402.967.4598.907.300.977.131.756.18171.10SD0.410.230.550.412.052.552.010.042.080.471.836.92Mean3.162.822.623.367.0877.576.451.117.231.986.25166.75SD0.240.220.600.891.9717.641.340.081.910.311.1713.74Mean2.762.632.823.527.6876.177.181.047.402.206.03162.15SD0.220.730.410.712.0718.941.520.091.210.410.835.77Mean2.042.412.443.486.0370.106.301.086.852.254.50169.73SD0.280.400.671.111.0710.471.080.181.800.500.565.25Mean1.881.623.043.864.8482.804.921.105.282.224.24156.64SD0.190.270.851.181.131.530.181.310.902.0220.88Mean3.102.892.562.536.480.006.201.026.282.105.33177.60SD1.291.031.261.492.020.002.060.082.101.331.953.14Mean1.481.594.665.767.830.006.901.258.603.855.23146.15SD0.490.382.132.641.690.001.540.232.141.330.9511.52Mean0.750.867.168.406.670.006.471.569.974.673.67141.47SD0.110.112.142.182.180.001.920.112.502.252.127.58Mean0.840.827.448.345.980.005.581.679.354.733.98133.58SD0.110.031.831.791.810.001.560.112.771.761.627.45Mean0.670.846.647.205.150.004.881.597.634.282.93135.95SD0.090.091.681.501.100.000.780.271.081.271.155.33Mean0.590.845.967.264.400.004.481.577.003.802.28133.15SD0.090.091.821.500.740.000.770.141.291.120.8215.14Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 299

PAGE 310

Appendix F (continued): Hemipristis elongata BOMean0.00SD0.00Mean27.23SD24.96Mean27.96SD30.22Mean42.62SD35.13Mean33.99SD3.47Mean34.30SD7.53Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 300 AARCARCACINALower, 1-24.113.8481.300.98172.73Lower, 4-52.962.7376.871.08164.45Lower 7-81.962.0176.451.09163.18Upper, 1-22.292.240.001.13161.88Upper, 4-50.790.840.001.62137.52Upper, 7-80.630.840.001.58134.55

PAGE 311

Appendix F (continued): Paragaleus pectoralis AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.730.843.183.332.600.003.051.003.051.382.30123.23SD0.350.140.660.660.200.000.370.030.310.290.146.88Mean1.560.723.954.002.800.003.321.083.581.602.45118.25SD0.320.110.440.470.240.000.290.100.100.200.298.80Mean1.311.964.284.408.70100.403.101.193.681.652.15121.38SD0.142.690.390.3812.200.360.070.250.170.134.18Mean1.260.544.154.532.2594.502.681.333.551.381.68123.83SD0.440.060.310.450.268.200.250.120.310.190.299.65Mean0.780.403.804.181.5373.681.781.943.351.150.83115.40SD0.390.090.670.730.3516.330.430.390.580.400.2916.27Mean0.380.343.433.601.1753.671.272.563.201.400.47106.00SD0.250.040.740.870.252.450.250.310.460.560.1510.37Mean1.320.762.452.931.8580.002.630.932.451.281.45148.53SD0.410.230.370.500.668.280.420.050.500.770.3011.45Mean0.870.454.004.032.6361.152.771.654.531.771.47118.97SD0.250.310.830.870.354.170.350.140.400.450.1512.17Mean0.850.634.434.702.7557.652.851.765.031.481.23107.63SD0.170.080.870.820.304.570.370.160.870.310.158.86Mean0.690.515.055.602.5854.902.701.905.131.751.20103.55SD0.090.040.600.720.459.170.410.150.830.340.183.86Mean0.630.554.584.802.5355.702.581.924.851.781.10105.25SD0.090.051.060.540.623.260.700.210.900.380.125.46Mean0.470.494.254.582.0853.202.082.164.451.700.80101.40SD0.040.060.540.290.385.510.320.180.370.080.0810.43Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 301

PAGE 312

Appendix F (continued): Paragaleus pectoralis BOMean27.99SD6.33Mean29.53SD13.30Mean28.82SD7.86Mean19.48SD8.40Mean13.37SD9.16Mean26.07SD11.57Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 302 AARCARCACINALower, 1-21.640.780.001.04120.74Lower, 4-51.291.2597.451.26122.60Lower 7-80.580.3763.672.25110.70Upper, 1-21.090.6170.581.29133.75Upper, 4-50.770.5756.281.83105.59Upper, 7-80.550.5254.452.04103.33

PAGE 313

Appendix F (continued): Hemigaleus microstoma AARCARBCWBWCHCADCLCIMCLNCWNHNAMean2.071.261.231.281.530.001.601.091.750.581.18151.68SD0.180.090.260.220.220.000.220.060.240.130.198.62Mean1.931.231.471.571.800.001.831.132.070.771.43141.63SD0.310.100.150.150.200.000.210.050.150.250.293.36Mean2.171.131.601.801.800.001.901.112.100.601.30145.40SDMean2.021.091.431.481.400.001.501.071.600.501.00149.53SD0.230.080.390.330.300.000.300.010.300.100.172.78Mean1.890.971.331.331.280.001.381.101.530.550.95148.80SD0.590.070.260.260.240.000.170.110.330.240.245.11Mean1.590.901.281.301.130.001.281.011.280.500.73147.28SD0.380.080.330.290.210.000.260.070.210.270.192.38Mean1.000.931.901.951.800.001.801.462.550.800.80133.75SD0.000.230.280.350.710.000.710.220.640.280.284.17Mean0.470.612.503.001.550.001.601.882.951.500.70127.05SD0.040.060.570.490.000.570.180.780.140.0010.39Mean0.530.613.253.252.000.002.001.793.551.400.75116.70SD0.100.050.640.640.570.000.570.120.780.140.212.97Mean0.450.662.832.971.870.001.871.733.201.230.57114.37SD0.110.050.550.490.420.000.420.110.560.150.214.71Mean0.220.592.903.001.700.001.501.982.951.150.25104.80SD0.020.050.000.000.140.000.140.230.070.210.0710.89Mean0.260.412.302.400.930.000.982.272.231.200.2896.33SD0.130.040.440.500.130.000.130.190.420.390.1010.01Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 303

PAGE 314

Appendix F (continued): Hemigaleus microstoma BOMean10.44SD6.19Mean-5.71SDMean3.94SD4.66Mean0.00SDMean11.59SDMean11.90SD10.38Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 304 AARCARCACINALower, 1-22.001.240.001.11146.65Lower, 4-52.091.110.001.09147.47Lower 7-81.740.930.001.05148.04Upper, 1-20.730.770.001.67130.40Upper, 4-50.490.630.001.76115.53Upper, 7-80.240.500.002.13100.56

PAGE 315

Appendix F (continued): Chlamydoselachus anguineus 305LoLoLoLoLoLoUUUUUU AARCARBCWBWCHCADCLCIMCLNCWNHNAMean2.631.163.403.403.9070.954.201.014.231.102.83153.13SD0.600.140.430.2026.380.260.040.310.170.3222.41Mean2.691.442.652.603.8044.903.951.014.001.052.80152.10SD0.580.120.210.004.670.070.050.140.070.4225.03Mean3.201.293.954.9061.205.301.035.401.103.60141.00SD0.590.212.051.842.260.071.980.281.5621.78Mean2.261.093.173.703.4057.903.771.023.831.102.47150.80SD0.180.160.580.200.350.030.380.170.253.30Mean2.301.063.102.903.3071.903.800.953.601.002.30161.80SDMean2.250.922.602.702.4061.502.900.902.600.801.80151.70SDMean3.531.792.673.504.6743.504.801.004.801.133.67152.33SD1.340.290.640.352.400.610.060.400.420.4223.20Mean2.631.154.334.404.9347.805.431.015.501.603.97134.03SD0.690.160.700.750.570.030.700.530.518.93Mean2.261.193.785.604.4048.904.801.044.883.263.50127.32SD1.150.141.281.1510.761.460.111.214.331.0113.62Mean3.001.184.504.605.3045.802.802.075.101.404.20133.50SD0.99Mean2.220.923.003.302.7558.603.101.073.300.902.00139.15SD0.790.260.000.7811.740.850.070.710.000.7111.38Mean1.830.973.503.703.4034.903.801.033.901.202.20154.60SDwer, 1wer, 2wer, 4wer, 5wer, 7wer, 8pper, 1pper, 8pper, 2pper, 4pper, 5pper, 7

PAGE 316

Appendix F (continued): Chlamydoselachus anguineus 306UULLLU BOMean-156.67SDMean-92.86SDMeanSDMean-202.53SDMean-68.63SDMean-45.71SDpper, 4-5pper, 7-8ower, 1-2ower, 4-5ower 7-8pper, 1-2 AARCARCACINALower, 1-22.661.3057.931.01152.62Lower, 4-52.731.1959.551.03145.90Lower 7-82.280.9966.700.92156.75Upper, 1-23.081.4745.651.01143.18Upper, 4-52.631.1847.351.55130.41Upper, 7-82.030.9446.751.05146.88

PAGE 317

Appendix F (continued): Hexanchus griseus AARCARBCWBWCHCADCLCIMCLNCWNHNAMean0.860.3723.1424.508.1347.6313.730.8612.074.273.6784.07SD0.120.066.867.113.626.224.020.134.741.461.332.00Mean0.590.3229.9030.839.7054.2017.731.1818.737.004.1583.80SD0.030.046.416.992.451.987.480.480.710.990.784.67Mean0.650.3025.8622.737.6252.8416.780.8013.084.422.8283.00SD0.100.048.105.572.422.915.660.385.621.521.063.94Mean0.550.2826.3026.946.2843.4013.580.8611.754.252.3379.93SD0.100.0410.079.371.6514.862.080.456.031.760.914.88Mean4.204.20SDMean2.703.00SDMean1.141.397.2011.088.54130.308.481.3611.305.205.36125.76SD0.300.613.294.612.25#DIV/0!2.830.334.372.501.8610.81Mean1.351.0611.2611.7010.0375.5311.051.1112.385.136.6387.70SD0.190.396.613.614.1910.045.040.155.732.362.4850.83Mean1.160.7011.4411.626.2074.756.951.168.053.604.05109.55SD0.130.054.033.933.543.614.030.014.601.701.485.16Mean1.230.7012.2612.826.0072.757.051.097.853.153.55100.20SD0.420.184.954.393.2512.233.890.074.741.480.492.26Mean0.610.4115.8516.755.5365.038.531.129.333.602.2092.23SD0.080.074.455.030.405.061.100.331.700.610.561.44Mean0.410.3212.4712.604.0776.576.571.147.473.531.47100.43SD0.170.102.402.291.7622.501.710.252.570.590.7521.43Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 307

PAGE 318

Appendix F (continued): Hexanchus griseus BOMean0.00SD0.00Mean-3.16SD9.02Mean-55.56SDMean-9.50SD24.68Mean-19.55SD25.80Mean5.10SD15.47Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 308 AARCARCACINALower, 1-20.730.3550.921.0283.93Lower, 4-50.600.2948.120.8381.46Lower 7-8Upper, 1-21.251.23102.921.24106.73Upper, 4-51.190.7073.751.13104.88Upper, 7-80.510.3670.801.1396.33

PAGE 319

Appendix F (continued): Notorhynchus cepidanus AARCARBCWBWCHCADCLCIMCLNCWNHNAMean0.670.3614.4315.785.5549.009.980.686.503.832.5082.88SD0.190.104.935.132.724.293.680.313.891.731.377.85Mean0.630.3417.0317.406.7746.6712.330.9711.905.173.2078.60SD0.130.036.045.200.855.851.190.090.720.650.266.56Mean0.590.3316.7016.785.7549.8310.200.959.884.902.8079.95SD0.090.076.416.052.736.733.640.113.902.181.076.58Mean0.370.3615.9016.185.6553.439.880.9710.08135.882.2875.63SD0.270.075.475.152.1311.292.690.294.46263.421.077.06Mean0.166.475.030.771.931.452.77SD0.094.591.650.150.590.811.43Mean0.193.904.250.702.150.962.05SD0.031.411.340.140.780.050.64Mean1.952.044.134.775.605.700.905.452.204.30170.90SD0.692.722.403.542.830.213.75Mean1.451.086.508.076.9059.405.971.458.373.604.90109.43SD0.280.203.463.053.3823.053.000.274.132.002.0120.36Mean1.210.6610.339.406.4376.576.631.419.233.504.13103.93SD0.100.125.065.072.454.413.670.084.981.822.048.83Mean1.110.719.1310.675.0570.105.101.527.754.404.90100.00SD0.045.034.773.613.390.005.16Mean0.630.4613.3012.956.0560.808.151.158.954.202.6584.95SD0.330.091.130.070.645.231.910.431.340.141.487.14Mean0.890.268.508.752.4561.705.050.773.754.504.0087.00SD0.094.815.302.052.900.091.77Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 309

PAGE 320

Appendix F (continued): Notorhynchus cepidanus BOMean-5.35SD4.66Mean-7.00SD6.74Mean0.00SD0.00Mean-44.81SD26.44Mean-16.39SD4.02Mean6.67SD9.43Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 310 AARCARCACINALower, 1-20.650.3547.830.8280.74Lower, 4-50.480.3451.630.9677.79Lower 7-80.171.21Upper, 1-21.701.5659.401.18140.17Upper, 4-51.160.6873.331.46101.97Upper, 7-80.760.3661.250.9685.98

PAGE 321

Appendix F (continued): Echinorhinus cookei AARCARBCWBWCHCADCLCIMCLNCWNHNAMean0.420.469.2711.104.2367.635.831.597.475.732.4392.43SD0.030.071.273.220.9512.672.391.154.551.020.5914.40Mean0.430.499.9311.134.8770.076.171.307.306.632.8799.50SD0.070.072.532.241.462.871.740.723.340.490.659.62Mean0.380.4210.7311.474.4362.436.531.638.507.002.6389.93SD0.050.062.582.830.865.353.201.084.811.250.153.04Mean0.420.3910.1311.333.8758.075.731.777.536.202.6086.83SD0.020.092.322.870.424.863.051.283.720.900.268.22Mean0.350.389.8310.533.7051.705.032.017.006.972.4077.33SD0.030.062.021.960.664.783.091.594.491.250.262.31Mean0.370.389.2310.173.5356.234.471.896.305.352.0780.83SD0.090.051.011.230.8411.422.291.403.860.350.326.91Mean0.490.4111.2512.134.4064.475.931.587.805.932.93102.90SD0.060.101.251.581.0412.972.380.993.250.700.5010.34Mean0.390.4011.6512.134.6865.35237.281.568.687.052.73100.40SD0.100.061.041.370.5714.96464.481.083.270.880.6016.49Mean0.470.4411.1311.954.8055.055.681.848.438.033.8594.25SD0.070.062.082.300.8210.872.621.023.432.911.8413.61Mean0.380.4610.98243.984.7753.136.271.507.336.732.60105.27SD0.060.012.34462.691.169.912.931.084.050.490.6225.71Mean0.380.4710.2811.704.7055.606.231.447.176.432.4787.23SD0.070.071.081.821.1323.512.711.013.490.060.4020.64Mean0.370.4110.3011.554.1345.876.201.457.576.272.3379.80SD0.040.062.162.001.4710.002.250.973.730.490.1210.41Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 311

PAGE 322

Appendix F (continued): Echinorhinus cookei BOMean-13.37SD18.90Mean-12.57SD13.15Mean-6.03SD10.44Mean-4.46SD5.30Mean-3.12SD6.04Mean-3.59SD7.18Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 312 AARCARCACINALower, 1-20.430.4768.851.4495.97Lower, 4-50.400.4160.251.7088.38Lower 7-80.360.3853.971.9579.08Upper, 1-20.440.4164.911.57101.65Upper, 4-50.430.4554.091.6799.76Upper, 7-80.380.4450.731.4583.52

PAGE 323

Appendix F (continued): Scymnodon ringens AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.861.994.003.657.6097.607.800.987.604.408.20131.20SD0.010.561.270.350.2815.840.710.050.280.851.568.91Mean1.791.565.254.708.1595.258.850.938.204.558.10136.30SD0.140.150.350.850.2121.140.070.010.000.350.008.91Mean1.691.424.804.357.40123.107.800.957.403.606.10143.10SD0.570.78Mean1.531.465.105.407.3084.507.200.966.954.206.45140.60SD0.070.370.710.710.854.810.850.071.340.140.4927.58Mean1.210.925.604.805.6084.705.601.025.704.305.20151.20SD0.710.57Mean1.191.025.255.255.25107.455.251.176.154.054.80133.45SD0.090.270.640.350.780.490.640.061.060.070.284.45Mean2.672.371.401.853.253.251.033.351.103.00162.20SD0.470.900.140.490.920.920.010.920.281.2724.47Mean2.532.241.651.803.753.901.094.201.403.55166.60SD0.040.440.210.141.201.410.071.270.571.4818.95Mean3.5316.282.152.2537.055.401.045.651.455.15165.10SD0.3119.410.210.0745.180.420.040.640.211.209.62Mean2.470.481.852.251.104.201.054.401.503.70142.10SD0.640.35Mean3.923.601.903.305.406.101.026.201.305.10167.90SD0.570.14Mean3.335.101.002.805.104.301.165.001.505.00163.70SDLower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 313

PAGE 324

Appendix F (continued): Scymnodon ringens BOMean29.29SD7.60Mean31.81SD4.58Mean28.91SD4.83Mean39.02SD55.19Mean0.00SD0.00Mean0.00SD0.00Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 314 AARCARCACINALower, 1-21.821.7796.430.95133.75Lower, 4-51.611.44103.800.95141.85Lower 7-81.200.9796.081.09142.33Upper, 1-22.602.310.001.06164.40Upper, 4-53.008.380.001.05153.60Upper, 7-83.634.350.001.09165.80

PAGE 325

Appendix F (continued): Somniosus microcephalus AARCARBCWBWCHCADCLCIMCLNCWNHNAMean0.370.466.106.372.7756.002.502.235.334.101.5389.93SD0.060.120.530.210.550.750.490.570.360.3512.72Mean0.380.366.376.872.2747.102.472.456.004.131.6077.30SD0.080.020.470.500.250.310.240.260.310.464.55Mean0.360.337.407.802.4334.302.402.566.134.831.7371.00SD0.080.020.360.350.210.100.070.210.150.403.98Mean0.370.307.077.472.1045.302.702.256.004.531.6780.10SD0.080.040.450.700.170.400.320.200.380.355.14Mean0.340.327.177.802.2750.602.133.406.474.401.5082.20SD0.060.030.310.660.210.721.650.510.260.205.81Mean0.300.337.257.202.4041.102.602.446.254.501.3572.10SD0.030.030.640.140.000.280.671.060.140.0711.31Mean1.561.662.603.134.434.371.164.772.103.33150.17SD0.150.410.440.681.681.330.370.720.691.4433.85Mean1.631.802.302.934.074.101.074.171.933.20148.10SD0.320.350.530.900.761.300.260.400.421.2319.87Mean1.551.672.473.474.074.031.154.532.033.13151.60SD0.120.500.720.911.271.400.121.160.841.2122.66Mean1.431.572.503.673.974.171.084.472.233.23158.10SD0.090.120.441.110.950.910.070.960.681.2114.92Mean1.041.342.903.433.353.551.003.551.901.95148.35SD0.260.290.700.670.070.070.000.070.140.350.21Mean1.441.862.873.205.304.201.134.702.453.60151.00SD0.290.370.320.261.840.850.130.420.491.4136.06Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 315

PAGE 326

Appendix F (continued): Somniosus microcephalus BOMean25.56SD7.03Mean21.86SD2.25Mean31.32SD2.25Mean-6.09SD43.36Mean-4.32SD33.72Mean-13.10SD22.68Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 316 AARCARCACINALower, 1-20.380.4151.552.3483.62Lower, 4-50.360.3139.802.4075.55Lower 7-80.320.3245.852.9277.15Upper, 1-21.591.730.001.12149.13Upper, 4-51.491.620.001.11154.85Upper, 7-81.241.600.001.07149.68

PAGE 327

Appendix F (continued): Aculeola nigra AARCARBCWBWCHCADCLCIMCLNCWNHNAMeanSDMean1.751.710.700.801.200.001.201.001.200.400.70167.60SDMean1.831.880.550.801.050.001.051.131.150.500.95176.10SD0.470.400.070.000.350.000.350.180.210.140.495.52Mean1.901.610.600.650.950.001.001.061.050.450.85146.45SD0.140.260.140.210.070.000.140.080.070.070.079.97Mean2.001.830.550.701.000.001.051.001.050.450.85167.50SD0.470.240.070.140.000.000.070.000.070.210.2117.68Mean2.331.330.600.700.800.000.801.130.900.300.70151.90SDMean1.671.330.600.700.800.000.901.000.900.300.50165.30SDMean1.751.540.500.750.750.000.801.060.850.350.60162.35SD0.350.290.140.070.070.000.000.090.070.070.0019.02MeanSDMean1.831.830.600.801.100.001.200.921.100.601.10179.20SDMeanSDMeanSDLower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 317

PAGE 328

Appendix F (continued): Aculeola nigra BOMeanSDMeanSDMeanSDMeanSDMeanSDMeanSDUpper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 318 AARCARCACINALower, 1-21.751.710.001.00167.60Lower, 4-51.871.750.001.09161.28Lower 7-82.171.580.001.06159.70Upper, 1-21.711.440.001.03163.83Upper, 4-51.831.830.000.92179.20Upper, 7-8

PAGE 329

Appendix F (continued): Etmopterus virens AARCARBCWBWCHCADCLCIMCLNCWNHNAMean0.241.540.771.270.6049.670.332.940.970.700.1771.10SD0.042.130.420.250.5218.840.060.630.150.200.0611.22Mean0.230.281.301.450.3555.950.403.071.200.900.2087.50SD0.040.040.420.490.0719.020.140.380.280.140.0019.37Mean0.320.371.101.200.4071.400.432.401.030.830.27101.07SD0.030.030.100.100.0028.280.060.170.060.120.0632.63Mean0.280.311.151.250.3557.450.452.281.000.750.2086.40SD0.080.080.070.070.075.300.070.670.140.210.007.50Mean0.250.331.201.250.4058.850.402.931.100.800.2084.80SD0.000.120.000.070.1417.320.141.040.000.000.0030.12Mean0.330.341.151.250.4057.400.452.351.050.750.2582.60SD0.060.100.070.210.1414.710.070.210.070.070.0712.87Mean1.030.660.630.750.4079.370.501.070.530.300.30128.37SD0.290.150.150.070.0027.800.000.230.120.100.1024.06Mean0.750.501.000.5085.600.701.000.700.400.30129.80SDMean1.330.830.600.800.5076.100.501.200.600.300.40140.10SDMeanSDMean1.670.830.600.700.5078.500.601.000.600.300.50135.20SDMeanSDLower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 319

PAGE 330

Appendix F (continued): Etmopterus virens BOMean23.38SD7.35Mean24.04SD1.36Mean34.78SDMeanSDMeanSDMeanSDUpper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 320 AARCARCACINALower, 1-20.230.9152.813.0179.30Lower, 4-50.300.3464.432.3493.73Lower 7-80.290.3458.132.6483.70Upper, 1-20.890.5882.481.03129.08Upper, 4-51.330.8376.101.20140.10Upper, 7-81.670.8378.50135.20

PAGE 331

Appendix F (continued): Etmopterus lucifer AARCARBCWBWCHCADCLCIMCLNCWNHNAMean0.330.321.631.870.5050.630.632.611.370.930.3077.50SD0.040.160.510.460.173.210.351.270.150.120.005.86Mean0.270.291.832.000.5363.770.602.971.731.270.3385.97SD0.110.040.310.300.156.120.170.570.310.230.156.11Mean0.360.291.932.200.5757.000.633.091.901.200.4381.87SD0.130.040.210.350.123.680.150.550.100.000.1512.06Mean0.260.292.032.270.6059.030.633.051.931.300.3388.33SD0.030.040.150.310.1010.240.060.190.250.100.0618.79Mean0.320.371.902.150.7060.900.702.791.951.400.45100.75SD0.010.050.140.210.143.540.140.060.350.280.0710.82Mean0.560.401.501.900.6059.600.702.001.400.900.5088.90SDMean1.881.071.031.171.1057.101.131.031.170.470.87143.93SD0.110.220.060.060.173.830.210.050.230.150.2510.42Mean1.631.191.001.101.1373.931.201.061.270.570.90147.37SD0.300.340.260.260.0614.350.000.130.150.120.0011.05Mean1.861.001.231.301.2357.901.301.081.400.571.03133.17SD0.480.140.060.000.155.270.100.010.100.060.157.86Mean1.900.921.201.401.1049.351.201.081.300.551.05112.40SD0.140.010.140.147.850.140.010.140.070.211.98Mean1.251.001.201.2051.701.400.931.300.801.00128.00SDMean1.000.751.201.200.9050.201.101.091.200.800.80119.20SDLower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 321

PAGE 332

Appendix F (continued): Etmopterus lucifer BOMeanSDMeanSDMeanSDMeanSDMeanSDMeanSDUpper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 322 AARCARCACINALower, 1-20.300.3157.202.7981.73Lower, 4-50.310.2958.023.0785.10Lower 7-80.440.3860.252.4094.83Upper, 1-21.751.1365.521.04145.65Upper, 4-51.880.9653.631.08122.78Upper, 7-81.130.8850.951.01123.60

PAGE 333

Appendix F (continued): Oxynotus bruniensis AARCARBCWBWCHCADCLCIMCLNCWNHNAMean0.620.7333.32.2111.32.51.082.72.11.3164.3SDMean0.580.72.32.41.6116.121.052.11.71166.9SDMean0.530.612.32.51.4951.51.672.50.9144.6SDMeanSDMeanSDMeanSDMean1.251.271.11.31.401.511.50.81158.8SDMean1.671.130.810.9011.11.10.60.7175SDMean0.90.9SDMeanSDMeanSDMeanSDLower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 323

PAGE 334

Appendix F (continued): Oxynotus bruniensis BOMean21.05SDMean64.00SDMeanSDMean17.39SDMeanSDMeanSDUpper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 324 AARCARCACINALower, 1-21.210.71113.701.07165.60Lower, 4-50.6195.001.67144.60Lower 7-8Upper, 1-20.601.200.001.05166.90Upper, 4-50.53Upper, 7-8

PAGE 335

Appendix F (continued): Dalatias licha AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.201.385.145.466.94109.006.541.076.964.805.65145.60SD0.200.240.750.640.6229.470.620.201.020.750.578.27Mean1.371.405.16111.587.05116.356.401.207.584.105.60143.25SD0.090.310.88235.580.668.410.780.200.610.280.0012.52Mean1.361.115.435.886.07109.156.531.056.874.906.65126.60SD0.050.080.320.920.708.980.400.070.570.140.0713.44Mean1.191.105.235.375.75116.406.351.036.504.355.10137.40SD0.210.070.120.320.493.820.780.170.280.640.149.19Mean1.090.954.935.104.6799.474.501.235.503.974.3393.47SD0.110.080.600.360.3516.210.520.160.260.150.5170.64Mean1.070.964.204.604.0389.873.671.455.273.433.67119.30SD0.020.070.200.100.155.350.150.451.420.120.1512.06Mean2.702.091.883.353.830.004.631.115.031.132.93164.70SD0.610.360.440.810.540.002.030.121.790.320.2117.90Mean2.152.731.983.104.680.004.431.155.051.603.33140.63SD0.601.380.680.340.540.000.570.140.410.300.4023.63Mean1.481.642.903.534.600.004.531.436.331.872.67131.83SD0.520.160.410.590.620.000.800.320.230.250.6124.44Mean1.321.472.853.603.970.003.871.325.131.872.47138.33SD0.230.420.440.320.610.000.460.201.030.250.575.83Mean1.221.053.103.633.230.003.331.605.231.932.20119.27SD0.540.270.200.650.810.000.670.340.810.550.6617.58Mean0.900.803.054.202.400.002.651.854.952.001.70107.80SD0.240.100.920.710.420.000.490.221.480.850.284.38Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 325

PAGE 336

Appendix F (continued): Dalatias licha BOMean18.87SD10.67Mean25.70SD6.03Mean21.32SD3.23Mean-12.12SD27.10Mean0.00SD0.00Mean4.44SD7.70Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 326 AARCARCACINALower, 1-21.281.39112.681.14144.43Lower, 4-51.271.11112.781.04132.00Lower 7-81.080.9694.671.34106.38Upper, 1-22.422.410.001.13152.67Upper, 4-51.401.550.001.38135.08Upper, 7-81.060.920.001.73113.53

PAGE 337

Appendix F (continued): Centrophorus granulosus AARCARBCWBWCHCADCLCIMCLNCWNHNAMean0.400.445.505.502.4085.252.801.714.803.151.25108.95SD0.000.030.420.000.001.340.140.010.280.210.0710.25Mean0.410.364.873.431.8083.172.431.603.932.901.17106.60SD0.050.041.212.300.6210.090.600.211.250.980.406.89Mean0.350.374.585.251.6590.182.031.984.002.951.03117.75SD0.060.070.690.350.136.740.130.260.420.530.199.42Mean0.430.384.685.181.7571.452.181.683.552.801.2095.75SD0.090.070.580.290.317.410.290.641.230.350.266.15Mean0.510.395.035.271.9780.302.371.884.402.701.3799.90SD0.120.100.450.400.495.730.450.130.560.170.329.58Mean0.290.284.304.601.2070.951.502.273.402.800.8096.30SD2.76Mean1.090.791.801.981.400.001.681.021.720.900.98145.76SD0.150.140.330.250.230.000.190.190.370.100.1814.64Mean1.080.871.702.101.430.001.601.051.670.931.00151.53SD0.230.220.400.280.230.000.260.130.230.060.1718.03Mean1.100.811.882.431.43140.201.631.131.830.900.95158.03SD0.290.330.540.520.460.420.080.410.350.3713.46Mean0.890.642.432.731.47127.001.901.052.001.100.93159.87SD0.280.200.720.550.060.300.090.350.260.1212.99Mean1.000.732.472.771.73137.402.030.921.561.331.27148.23SD0.430.320.500.400.6826.300.670.711.110.250.4520.10Mean0.990.772.553.002.00118.002.401.062.451.601.60139.39SD0.270.080.640.570.710.990.170.640.140.578.37Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 327

PAGE 338

Appendix F (continued): Centrophorus granulosus BOMeanSDMeanSDMeanSDMeanSDMeanSDMeanSDUpper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 328 AARCARCACINALower, 1-20.400.4084.211.66107.78Lower, 4-50.390.3780.811.83106.75Lower 7-80.400.3475.632.0798.10Upper, 1-21.080.83-9999.001.04148.65Upper, 4-51.000.73133.601.09158.95Upper, 7-80.990.75127.700.99143.81

PAGE 339

Appendix F (continued): Squalus acanthias AARCARBCWBWCHCADCLCIMCLNCWNHNAMean0.480.442.342.441.0268.861.141.952.221.320.62108.58SD0.100.090.330.350.1312.140.050.300.330.220.0416.32Mean0.400.367.662.721.1272.621.162.112.421.560.60110.38SD0.110.1811.370.280.0819.990.110.370.270.230.1013.49Mean0.380.392.902.981.1466.741.222.282.761.920.72110.94SD0.090.060.250.300.2117.230.180.360.380.320.1611.96Mean0.340.352.903.041.0068.781.122.502.701.880.64104.52SD0.110.090.200.340.2517.350.250.580.280.040.2115.09Mean0.360.352.903.061.0071.561.082.562.701.880.64102.34SD0.110.060.210.320.169.870.160.590.360.410.1114.20Mean0.340.342.722.840.9277.751.042.402.481.840.62107.04SD0.060.040.110.090.133.870.110.210.200.150.1310.53Mean0.540.471.832.000.8589.851.131.441.481.250.65121.23SD0.140.090.260.340.1012.850.340.600.370.250.0617.11Mean0.600.481.902.050.9079.801.181.431.551.100.65127.50SD0.090.110.080.170.1810.260.360.490.240.220.1317.34Mean0.530.462.062.240.9480.581.061.791.841.200.62118.12SD0.080.090.220.270.1713.710.190.380.170.190.0416.39Mean0.510.432.152.300.9387.731.081.781.851.180.60119.93SD0.070.080.190.080.2113.050.260.360.210.190.1412.53Mean0.460.402.152.280.8580.400.952.131.931.280.58107.20SD0.090.100.210.130.2416.500.260.490.100.170.1014.47Mean0.370.392.202.340.8671.201.221.881.761.450.56107.96SD0.150.100.230.230.2113.540.661.060.440.210.1515.40Lower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 329

PAGE 340

Appendix F (continued): Squalus acanthias BOMean22.43SD1.19Mean20.43SD3.35Mean19.14SD6.31Mean18.61SD2.52Mean19.55SD2.19Mean18.06SD4.40Upper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 330 AARCARCACINALower, 1-20.440.4070.742.03109.48Lower, 4-50.360.3767.762.39107.73Lower 7-80.350.3474.662.48104.69Upper, 1-20.570.4784.831.44124.36Upper, 4-50.520.4584.151.78119.02Upper, 7-80.410.3975.802.00107.58

PAGE 341

Appendix F (continued): Squatina dumeril AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.621.632.404.303.900.004.800.944.502.103.40147.30SDMean1.250.974.556.253.700.006.001.006.054.354.30144.05SD0.720.652.192.050.850.001.700.031.913.180.855.16Mean1.270.844.706.903.900.004.301.195.152.603.30129.05SD0.120.851.410.140.000.140.170.925.73Mean1.360.835.207.404.050.004.451.185.252.253.00124.20SD0.160.242.121.560.490.000.920.031.200.780.714.38Mean1.500.704.605.803.200.004.500.803.601.602.40137.60SDMean1.450.646.107.003.900.004.201.385.802.002.90113.60SDMean1.291.113.705.003.500.003.651.234.401.552.00151.90SD0.020.522.262.400.570.001.200.170.850.350.421.84Mean1.330.914.254.804.300.004.601.155.302.403.20131.50SD0.640.85Mean1.090.933.956.453.500.003.501.454.902.302.55125.00SD0.200.241.480.640.420.000.990.370.140.420.9210.75Mean1.530.674.906.353.900.004.401.205.301.902.90128.90SD1.270.21Mean1.450.874.256.553.350.004.101.003.901.752.50135.70SD0.170.342.052.330.350.001.410.280.280.490.429.62Mean1.710.685.006.203.400.004.201.054.401.402.40128.70SDLower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 331

PAGE 342

Appendix F (continued): Squatina dumeril BOMean-21.98SDMean-26.55SD6.50MeanSDMean-10.67SD15.08Mean-51.16SD12.08Mean-9.72SDUpper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 332 AARCARCACINALower, 1-21.441.300.000.97145.68Lower, 4-51.310.830.001.19126.63Lower 7-81.480.670.001.09125.60Upper, 1-21.311.010.001.19141.70Upper, 4-51.310.800.001.33126.95Upper, 7-81.580.770.001.02132.20

PAGE 343

Appendix F (continued): Pristiophorus cirratum AARCARBCWBWCHCADCLCIMCLNCWNHNAMean1.20.52.83.21.402.30.831.911.2134.3SDMean0.890.382.42.50.901.41.291.80.90.8108.3SDMean0.90.482.52.81.201.71.18210.9123.1SDMean1.140.462.42.71.101.41.361.90.70.8125.1SDMean1.20.441.81.90.801.21.081.30.50.6138.2SDMeanSDMean1.080.583.13.61.802.40.962.31.21.3136.5SDMean10.552.22.51.201.41.341.80.90.9138.6SDMean0.910.672.32.71.301.71.121.91.11135.4SDMean1.220.592.73.21.6021.152.30.91.1131.7SDMean10.631.92.21.201.41.211.70.70.7136SDMean0.860.4522.30.901.11.551.70.70.6132.6SDLower, 1Lower, 2Lower, 4Lower, 5Lower, 7Lower, 8Upper, 1Upper, 8Upper, 2Upper, 4Upper, 5Upper, 7 333

PAGE 344

Appendix F (continued): Pristiophorus cirratum BOMean14.04SDMean10.91SDMeanSDMean0SDMean0SDMean0SDUpper, 4-5Upper, 7-8Lower, 1-2Lower, 4-5Lower 7-8Upper, 1-2 334 AARCARCACINALower, 1-21.040.790.001.06121.30Lower, 4-51.020.950.001.27124.10Lower 7-81.200.001.08138.20Upper, 1-21.040.800.001.15137.55Upper, 4-51.071.230.001.13133.55Upper, 7-80.931.000.001.38134.30

PAGE 345

Appendix G: Significant Habitat Variables from Evolutionary Analysis 335

PAGE 346

Appendix G: Abbreviations for Shape and Model can be found in Chapter 3. Subheadings under Variable are dummy variables of Habitat and Depth: BP= benthopelgic/not; P=pelgaic/not; O=oceanic/not; B=bathyal/not; CB=costal-bathyal/not; CO=costal-oceanic/not. A = anterior; P = posterior; Bold type = lowest AIC; = p<0.05. ToothShape ModelAICVariableSlopedfPartial-Ftp2, UpperAAROLS109.06Habit2, 351.110.341BP-0.9635-1.400.170P-0.6735-1.010.320Depth4, 350.790.540O0.3135-0.620.539B0.21350.290.774CB0.77354.44>0.001*CO0.54350.970.339PGLS105.07Habit2, 351.170.322BP0.5935-0.630.533P0.53350.220.827Depth4, 351.240.312O0.45351.170.250B0.64350.440.663CB0.55352.070.046*CO0.56351.340.1895, UpperAAROLS67.94Habit2,320.450.642BP-0.4032-0.900.375P0.4332-0.940.354Depth4,322.140.099O0.32320.680.501B0.4732-0.190.851CB0.36322.140.040*CO0.38321.630.113PGLS76.88Habit2,320.010.990BP0.01320.040.968P0.04320.090.929Depth4,320.860.498O0.28320.810.424B-0.0632-0.130.987CB0.72321.610.117CO0.45321.020.315 336

PAGE 347

Appendix G (continued): ToothShape ModelAICVariableSlopedfPartial-FtpLower, 1CAROLS120.50Habit2, 350.480.623BP-0.7735-0.980.334P-0.6335-0.830.412Depth4, 352.970.033*O0.13350.230.820B2.86353.410.002*CB0.75350.020.984CO-0.0835-0.120.905PGLS128.59Habit2, 350.040.961BP-0.1235-0.020.984P-0.1735-0.250.804Depth4, 353.560.015*O0.51350.860.396B3.00353.540.001*CB0.95351.300.202CO0.34350.450.656Upper, 7CIOLS30.56Habit2,320.730.490BP0.00320.120.905P0.19320.730.471Depth4,322.830.037*O0.0032-0.040.968B0.03320.900.375CB-0.0432-1.880.069CO-0.2732-1.100.280PGLS41.73Habit2,320.340.714BP-0.2332-0.080.937P-0.1132-0.450.656Depth4,321.050.397O0.10320.470.642B0.19320.630.533CB-0.2032-0.740.465CO-0.0132-0.290.774 337

PAGE 348

Appendix G (continued): ToothShape ModelAICVariableSlopedfPartial-FtpUpper, 5NAOLS398.99Habit2,320.010.990BP0.28320.001.000P-1.5132-0.010.992Depth4,321.750.163O-5.9132-0.260.797B8.27322.500.018*CB-0.6732-0.010.992CO9.76320.670.508PGLS414.95Habit2,320.070.933BP-7.7132-0.220.827P0.15320.001.000Depth4,321.480.231O20.2532-0.770.447B70.45321.910.065CB-19.5232-0.570.573CO-12.0032-0.360.720Upper, 7NAOLS398.75Habit2,324.030.028*BP-80.7432-2.560.015*P-84.7732-2.830.008*Depth4,322.730.046*O0.94320.001.000B-106.3432-3.22<0.001*CB10.40320.430.670CO9.34320.340.736PGLS413.22Habit2,322.120.137BP-67.7632-1.980.056P-56.6832-1.910.650Depth4,322.740.046*O-29.9832-1.170.251B-115.6032-3.21.0.003*CB-19.1832-0.610.546CO-32.5432-1.000.325 338

PAGE 349

Appendix G (continued): ToothShape ModelAICVariableSlopedfPartial-FtpLower, 7-8BOOLS324.86Habit2,280.430.655BP1.52280.080.937P10.88280.580.567Depth4,281.140.360O9.90280.700.490B7.70280.380.707CB-9.9928-0.640.528CO-3.1028-0.170.866PGLS311.14Habit2,280.700.505BP-17.9428-1.110.276P-46.9028-0.380.707Depth4,282.850.042*O3.39280.330.074B-1.3028-0.010.992CB-23.4728-1.760.089CO-16.5228-1.140.264Upper, 4-5BOOLS292.46Habit2,273.470.046*BP21.99271.600.121P32.12272.440.022*Depth4,273.900.130*O7.82270.790.436B34.64272.400.024*CB-15.8327-1.380.179CO-5.6827-0.440.663PGLS291.73Habit2,275.210.012*BP-2.2527-0.180.858P21.63271.990.057Depth4,272.070.113O0.23270.020.984B23.18271.800.083CB-14.7527-1.280.211CO-4.5527-0.360.722 339

PAGE 350

Appendix G (continued): ToothShape ModelAICVariableSlopedfPartial-FtpA, LowerAAROLS133.54Habit2,360.500.611BP-0.8936-1.000.324P-0.7336-0.850.400Depth4,361.410.250O0.34360.530.600B2.15362.270.029*CB0.10360.150.882CO0.19360.260.796PGLS132.21Habit2,360.090.914BP-0.2836-0.360.721P-0.2936-0.410.684Depth4,362.520.058O0.71361.180.246B2.21362.580.014CB1.483620.30<0.001*CO1.19361.590.121A, LowerCAROLS107.33Habit2,360.620.544BP-0.7336-1.120.270P-0.6336-0.990.329Depth4,361.520.217O0.07360.150.882B1.69362.420.021*CB-0.0536-0.100.921CO-0.10360.180.858PGLS113.11Habit2,360.030.971BP-0.1536-0.230.819P-0.1036-0.170.866Depth4,362.130.097O0.35360.730.470B1.92362.800.008*CB0.65361.110.274CO0.50360.830.412 340

PAGE 351

Appendix G (continued): ToothShape ModelAICVariableSlopedfPartial-FtpP, LowerCAROLS668.84Habit2,340.930.404BP28.82340.440.663P-14.4434-0.230.820Depth4,341.860.140O23.96340.010.992B-0.19340.001.000CB144.16340.280.781CO-905.3834-1.670.104PGLS675.34Habit2,342.830.073BP828.65341.260.216P-138.6834-0.240.812Depth4,343.580.015*O354.36340.720.476B190.62340.270.789CB37.73340.010.992CO-1013.7234-1.640.110P, UpperNAOLS377.19Habit2,322.020.015BP-42.4632-1.770.086P-45.6232-2.000.054Depth4,321.800.153O1.44320.010.992B-65.2432-2.610.014*CB5.40320.290.774CO9.95320.470.612PGLS388.40Habit2,321.070.355BP-36.4032-1.460.154P-25.6732-1.190.243Depth4,321.670.181O-21.0032-1.120.271B-66.1132-2.530.017*CB-20.6632-0.900.375CO-25.8132-1.090.284 341

PAGE 352

Appendix H: Significant Diet Variables from Evolutionary Analysis 342

PAGE 353

Appendix H: Abbreviations for Shape, Variable, and Model can be found in Chapter 3. A = anterior; L = lateral; P = posterior; Bold type = lowest AIC; = p<0.05. ToothShape dfModelAICVariableSlopePartial-Fp5, UpperAAR1,29OLS79.71Elasmobranch0.110.180.675Teleost-0.120.660.427Shrimp-0.100.130.721Crab0.040.020.889Cephalopod0.210.980.330Worm0.210.190.666Hard Ech/Moll0.220.190.666Mammal0.040.020.889Turtle0.501.070.310PGLS79.87Elasmobranch0.140.260.614Teleost0.120.140.711Shrimp-0.010.001.000Crab0.080.090.766Cephalopod0.444.870.035*Worm0.350.770.387Hard Ech/Moll0.430.960.335Mammal0.492.660.114Turtle0.100.050.8258, UpperCAR1,32OLS442.67Elasmobranch-31.213.000.093Teleost-6.560.140.711Shrimp-13.070.510.480Crab9.000.230.635Cephalopod-9.130.400.532Worm-70.784.240.048*Hard Ech/Moll-34.731.020.320Mammal19.790.750.393Turtle-4.730.020.888PGLS446.04Elasmobranch-30.853.600.067Teleost-2.860.030.864Shrimp7.000.150.701Crab0.880.001.000Cephalopod-2.910.040.843Worm-64.804.940.034*Hard Ech/Moll-34.331.140.294Mammal46.894.500.042*Turtle-36.801.470.234 343

PAGE 354

Appendix H (continued): ToothShape dfModelAICVariableSlopePartial-Fp1, UpperCA1,31OLS438.58Elasmobranch-40.594.220.049*Teleost28.182.360.135Shrimp-31.672.590.118Crab11.760.560.460Cephalopod-18.801.470.235Worm-48.611.760.194Hard Ech/Moll-25.830.510.481Mammal7.580.080.779Turtle-8.770.060.808PGLS453.97Elasmobranch-32.552.480.126Teleost22.371.150.292Shrimp-2.800.010.921Crab-9.080.200.658Cephalopod-12.120.410.527Worm-13.790.140.711Hard Ech/Moll-39.830.970.332Mammal7.220.070.793Turtle-13.880.130.7212, UpperCA1,31OLS423.09Elasmobranch-33.604.250.048*Teleost28.533.530.070Shrimp-25.632.230.146Crab0.660.001.000Cephalopod-22.803.160.085Worm-3.670.010.921Hard Ech/Moll-14.980.250.621Mammal2.970.020.889Turtle-7.630.060.808PGLS438.60Elasmobranch-25.792.240.145Teleost29.462.890.099Shrimp-10.470.310.582Crab-1.450.010.921Cephalopod-19.001.620.213Worm32.900.720.403Hard Ech/Moll-10.220.090.766Mammal10.090.200.658Turtle-22.150.490.489 344

PAGE 355

Appendix H (continued): ToothShape dfModelAICVariableSlopePartial-Fp5, UpperCA1, 27OLS391.39Elasmobranch-24.071.560.222Teleost15.220.560.461Shrimp-11.120.240.628Crab-0.860.001.000Cephalopod-17.151.200.283Worm-100.534.430.045*Hard Ech/Moll-45.221.630.213Mammal1.050.001.000Turtle-9.130.070.793PGLS403.97Elasmobranch-1.170.001.000Teleost20.400.730.400Shrimp16.130.500.486Crab-13.150.380.543Cephalopod2.340.020.889Worm-56.941.720.201Hard Ech/Moll-36.510.950.338Mammal7.840.090.767Turtle-30.230.760.3911, LowerCI1,31OLS68.90Elasmobranch0.211.000.325Teleost0.020.010.921Shrimp0.030.020.889Crab0.160.520.476Cephalopod-0.191.150.292Worm-0.110.070.793Hard Ech/Moll0.100.050.825Mammal-0.700.060.808Turtle-0.230.320.576PGLS46.07Elasmobranch0.060.160.692Teleost0.100.440.512Shrimp0.050.110.742Crab0.150.920.345Cephalopod-0.264.470.043*Worm-0.321.570.220Hard Ech/Moll-0.150.300.588Mammal-0.271.950.173Turtle-0.150.350.558 345

PAGE 356

Appendix H (continued): ToothShape dfModelAICVariableSlopePartial-Fp5, UpperCI1,29OLS5.10Elasmobranch-0.131.770.194Teleost0.6530.58<0.001*Shrimp-0.121.260.271Crab0.141.820.188Cephalopod-0.050.330.570Worm0.120.380.542Hard Ech/Moll0.394.120.052Mammal0.182.040.164Turtle-0.140.540.469PGLS10.93Elasmobranch-0.040.210.650Teleost0.5115.17<0.001*Shrimp-0.255.690.024*Crab0.276.890.014*Cephalopod-0.070.710.406Worm0.090.310.582Hard Ech/Moll0.497.310.011*Mammal0.192.390.133Turtle-0.100.340.5647, UpperCI1,29OLS34.94Elasmobranch-0.181.540.225Teleost0.262.890.100Shrimp-0.385.000.033*Crab0.000.010.210Cephalopod-0.070.300.588Worm0.421.280.267Hard Ech/Moll0.513.140.087Mammal0.261.940.174Turtle-0.220.630.434PGLS35.85Elasmobranch-0.141.280.267Teleost0.110.630.434Shrimp-0.4811.00.003*Crab0.090.360.553Cephalopod-0.151.630.212Worm0.371.300.264Hard Ech/Moll0.514.100.052Mammal0.090.280.601Turtle0.010.001.000 346

PAGE 357

Appendix H (continued): ToothShape dfModelAICVariableSlopePartial-Fp8, UpperCI1,27OLS51.96Elasmobranch-0.160.690.413Teleost0.413.970.057Shrimp-0.597.170.012*Crab0.010.001.000Cephalopod-0.181.240.275Worm0.320.470.499Hard Ech/Moll0.764.240.049*Mammal0.210.800.038Turtle-0.441.590.218PGLS68.22Elasmobranch-0.160.630.434Teleost0.261.010.324Shrimp-0.647.610.012*Crab-0.070.010.921Cephalopod-0.282.120.157Worm0.350.410.527Hard Ech/Moll0.622.430.131Mammal-0.060.050.825Turtle-0.300.650.4274, UpperNA1, 32OLS367.54Elasmobranch21.288.350.007*Teleost-3.260.220.642Shrimp6.700.770.387Crab0.790.010.920Cephalopod15.786.970.013*Worm-7.390.280.600Hard Ech/Moll-30.144.660.039*Mammal-7.820.690.412Turtle-6.770.240.628PGLS378.17Elasmobranch14.814.170.049*Teleost-1.920.060.808Shrimp14.043.050.903Crab-4.630.380.542Cephalopod14.905.400.027*Worm-4.990.150.701Hard Ech/Moll-36.886.50.016*Mammal-6.360.420.522Turtle-6.160.210.650 347

PAGE 358

Appendix H (continued): ToothShape dfModelAICVariableSlopePartial-Fp7, UpperNA1,29OLS411.89Elasmobranch38.974.600.041*Teleost-30.382.510.124Shrimp27.251.650.209Crab1.640.010.921Cephalopod3.230.040.843Worm-12.160.070.793Hard Ech/Moll-33.370.860.361Mammal-0.070.001.000Turtle-8.030.060.808PGLS421.62Elasmobranch41.435.450.027*Teleost-17.700.790.381Shrimp38.093.520.071Crab-10.590.270.607Cephalopod13.090.660.423Worm-4.770.010.921Hard Ech/Moll-37.861.130.297Mammal0.130.001.000Turtle-22.650.470.4988, UpperNA1,27OLS361.64Elasmobranch17.171.900.179Teleost-31.265.270.030*Shrimp12.890.800.379Crab-12.250.840.368Cephalopod10.971.060.312Worm13.790.200.658Hard Ech/Moll-33.431.910.178Mammal-12.070.600.445Turtle6.290.080.780PGLS373.10Elasmobranch10.730.720.404Teleost-33.144.540.042*Shrimp9.260.410.527Crab-9.790.480.494Cephalopod12.931.200.283Worm13.190.150.702Hard Ech/Moll-28.251.340.257Mammal-5.390.110.723Turtle-0.650.001.000 348

PAGE 359

Appendix H (continued): ToothShape dfModelAICVariableSlopePartial-Fp4-5, LowerBO1,28OLS357.83Elasmobranch9.450.830.370Teleost-9.170.750.394Shrimp-2.330.040.843Crab-14.361.650.210Cephalopod5.940.460.503Worm2.390.020.889Hard Ech/Moll3.410.030.864Mammal-0.360.001.000Turtle-0.900.001.000PGLS342.98Elasmobranch-1.520.040.843Teleost-17.294.350.046*Shrimp-14.953.180.085Crab-5.630.470.499Cephalopod-6.781.040.317Worm-8.450.440.513Hard Ech/Moll-3.430.060.808Mammal-8.390.740.397Turtle13.861.080.3084-5, UpperBO1,24OLS312.48Elasmobranch3.120.110.743Teleost2.860.070.794Shrimp-4.060.140.712Crab-4.620.210.651Cephalopod1.860.050.825Worm-4.400.060.809Hard Ech/Moll-5.880.110.743Mammal7.190.370.589Turtle-14.970.770.389PGLS294.45Elasmobranch-5.350.720.405Teleost-7.840.940.342Shrimp-24.2110.560.003*Crab6.460.010.921Cephalopod-6.941.390.250Worm0.040.001.000Hard Ech/Moll5.290.180.675Mammal-11.281.840.188Turtle10.250.080.780 349

PAGE 360

Appendix H (continued): ToothShape dfModelAICVariableSlopePartial-Fp7-8, UpperBO1,25OLS333.34Elasmobranch5.070.220.643Teleost-15.881.660.209Shrimp-1.340.010.921Crab-6.630.300.589Cephalopod-0.070.001.000Worm-9.560.120.732Hard Ech/Moll-8.190.150.702Mammal2.130.020.889Turtle-8.400.170.684PGLS317.91Elasmobranch-6.510.710.407Teleost-22.415.580.026*Shrimp-16.012.830.105Crab-6.780.610.442Cephalopod-10.662.330.140Worm6.180.100.755Hard Ech/Moll-14.560.940.342Mammal-12.701.590.219Turtle19.642.030.1671, UpperAAR1,39OLS120.25Soft-0.100.110.742Medium0.210.390.540Hard0.634.290.045*PGLS121.54Soft0.040.030.860Medium0.470.290.590Hard0.042.040.1605, UpperCI1,35OLS10.33Soft0.010.010.920Medium0.4716.75<0.001*Hard0.225.980.020*PGLS16.68Soft-0.101.500.230Medium0.348.560.006*Hard0.152.540.1201-2, UpperBO1,33OLS369.95Soft-0.440.001.000Medium-28.554.470.042*Hard-20.683.070.090PGLS371.42Soft-10.171.070.309Medium-31.646.170.019*Hard-21.933.580.067 350

PAGE 361

Appendix H (continued): ToothShape dfModelAICVariableSlopePartial-FpP, UpperCAR1,29OLS92.62Elasmobranch0.250.710.406Teleost-0.160.240.628Shrimp-0.190.290.594Crab-0.170.290.594Cephalopod0.382.180.151Worm0.750.950.338Hard Ech/Moll0.550.850.364Mammal0.090.050.825Turtle0.110.020.889PGLS92.02Elasmobranch0.170.440.512Teleost-0.140.230.635Shrimp-0.050.030.864Crab-0.361.400.246Cephalopod0.484.110.052Worm0.781.360.253Hard Ech/Moll0.400.580.453Mammal0.774.410.045*Turtle-0.871.550.223A, LowerCA1,32OLS440.07Elasmobranch-30.733.230.082Teleost-6.470.140.711Shrimp-14.610.680.416Crab9.640.300.589Cephalopod-9.170.430.517Worm-73.644.950.033*Hard Ech/Moll-31.930.920.345Mammal10.590.210.650Turtle42.970.900.350PGLS445.49Elasmobranch-34.924.750.037*Teleost-3.170.030.864Shrimp4.990.070.793Crab-0.650.001.000Cephalopod-5.060.120.731Worm-65.375.060.032*Hard Ech/Moll-32.161.010.322Mammal39.002.850.101Turtle-70.010.030.864 351

PAGE 362

Appendix H (continued): ToothShape dfModelAICVariableSlopePartial-FpA, UpperCA1,32OLS437.89Elasmobranch-41.125.990.020*Teleost28.863.230.082Shrimp-32.643.630.066Crab6.900.170.683Cephalopod-19.862.190.149Worm-47.382.210.147Hard Ech/Moll-18.530.340.564Mammal-6.080.070.793Turtle45.591.080.607PGLS456.51Elasmobranch-37.784.140.050Teleost26.841.970.170Shrimp-13.400.400.532Crab-5.830.010.921Cephalopod-19.931.500.230Worm-21.660.440.512Hard Ech/Moll-29.740.660.423Mammal-1.290.001.000Turtle23.700.220.642A, UpperCI1,32OLS439.86Elasmobranch-16.040.880.355Teleost-18.981.220.278Shrimp-8.170.210.650Crab7.400.180.674Cephalopod16.761.450.237Worm3.890.040.843Hard Ech/Moll-34.891.100.302Mammal52.405.040.032Turtle-29.690.430.517PGLS447.42Elasmobranch-14.610.790.381Teleost-23.051.790.190Shrimp-16.900.800.378Crab2.620.020.888Cephalopod23.372.540.121Worm25.750.750.293Hard Ech/Moll-4.640.020.888Mammal42.913.290.079Turtle-19.350.190.666 352

PAGE 363

Appendix H (continued): ToothShape dfModelAICVariableSlopePartial-FpA, UpperNA1,33OLS402.50Elasmobranch20.884.530.041*Teleost-13.922.210.147Shrimp10.481.090.304Crab5.030.260.614Cephalopod15.073.680.064Worm-2.470.020.888Hard Ech/Moll-26.892.090.158Mammal3.400.060.808Turtle-24.370.880.355PGLS421.72Elasmobranch15.482.050.162Teleost-12.741.280.266Shrimp3.860.100.754Crab4.120.130.721Cephalopod19.704.190.049*Worm-1.890.010.921Hard Ech/Moll-16.680.600.444Mammal4.430.080.779Turtle-4.220.020.888L, UpperNA1,33OLS432.27Elasmobranch21.742.510.123Teleost-27.634.450.043*Shrimp-1.670.010.921Crab-1.280.010.921Cephalopod10.200.860.361Worm8.490.100.754Hard Ech/Moll-29.711.310.261Mammal-2.150.010.921Turtle-24.530.470.498PGLS443.33Elasmobranch19.742.020.165Teleost-29.864.240.047Shrimp9.230.340.564Crab-8.910.380.542Cephalopod5.250.180.674Worm1.540.001.000Hard Ech/Moll-46.462.810.103Mammal-8.330.170.683Turtle-28.920.580.452 353

PAGE 364

Appendix H (continued): ToothShape dfModelAICVariableSlopePartial-FpP, UpperNA1,29OLS377.93Elasmobranch30.847.150.012*Teleost28.635.320.028Shrimp22.472.700.111Crab-4.350.200.658Cephalopod8.150.680.416Worm0.490.001.000Hard Ech/Moll-33.152.040.164Mammal0.780.001.000Turtle-32.061.120.299PGLS385.17Elasmobranch30.987.900.009*Teleost-23.463.490.072Shrimp32.996.290.018*Crab-17.761.830.190Cephalopod16.092.540.122Worm13.400.220.643Hard Ech/Moll-35.102.460.128Mammal8.810.310.582Turtle-58.813.830.060L, UpperCI1,38OLS61.55Soft-0.161.180.284Medium0.050.010.921Hard0.191.400.244PGLS50.78Soft-0.141.530.224Medium0.171.470.233Hard0.305.160.029* 354

PAGE 365

About the Author Born in Park Ridge, IL on May 9, 1977, Lisa Beth Whitenack was interested in fossils and fishes from an early age. After gra duating from Schaumburg High School in 1995, Lisa attended the University of Illinois at Urbana-Champaign, receiving her B.S. in Geology in 1999. She earned a M.S. in Geol ogical Sciences from Michigan State University in 2001, then joined Dr. Philip Mott as lab to earn her doctorate in Biology on the biomechanics and evolution of shark teeth. She has taught a variety of classes from geology to biology, and earned the Provost s Commendation for Outstanding Teaching by a Graduate Teaching Assistant, Biology Department Outstanding Graduate Teaching Assistant award, and Jeffrey C. and Carol A. Carrier Award for the Outstanding Student Poster Presentation. Upon completion of her doctorate, Lisa has authored five papers and will join the Department of Geology at the Un iversity of South Flor ida as a postdoctoral fellow.