The Evolution Of Teeth: A Review And Phylogenetic Analysis Using Categorical Data

Item Type text; Electronic Thesis

Authors Lee, Davis Min

Publisher The University of Arizona.

Rights Copyright © is held by the author. Digital access to this material is made possible by the University Libraries, University of Arizona. Further transmission, reproduction or presentation (such as public display or performance) of protected items is prohibited except with permission of the author.

Download date 28/09/2021 19:42:44

Item License http://rightsstatements.org/vocab/InC/1.0/

Link to Item http://hdl.handle.net/10150/632849

THE EVOLUTION OF VERTEBRATE TEETH:

A REVIEW AND PHYLOGENETIC ANALYSIS USING CATEGORICAL DATA

By

DAVIS MIN LEE

______

A Thesis Submitted to The Honors College

In Partial Fulfillment of the Bachelors degree With Honors in

Biology

THE UNIVERSITY OF ARIZONA

M A Y 2 0 1 9

Approved by:

______

Dr. John Wiens Department of Ecology and Evolutionary Biology

Lee 1

Table of Contents

ABSTRACT ...... 3

INTRODUCTION ...... 4

MATERIALS AND METHODS ...... 6

RESULTS ...... 9

LOCATION ...... 9 REPLACEMENT ...... 11 ATTACHMENT ...... 12

DISCUSSION ...... 14

CONCLUSION ...... 19

REFERENCES ...... 20

APPENDIX 1: CHARACTER MAPS ...... 26

FIGURES 1-11: LOCATION ...... 26 FIGURES 12-15: REPLACEMENT ...... 32 FIGURES 16-21: ATTACHMENT ...... 34

APPENDIX 2: SUPPLEMENTAL TABLES ...... 37

SUPPLEMENTAL TABLES 1.1-1.3: LOCATION ...... 37 SUPPLEMENTAL TABLES 2.1-2.3: REPLACEMENT AND ATTACHMENT ...... 40

2

Lee 3

Abstract

The crucial importance in phylogenetic analyses lies in their ability to detail the evolution of living by comparing homologous traits that arose over time. Through them, these analyses not only tell what was derived from what, but also reveal how environmental circumstances drove the formation of characteristics favorable for that time. Though teeth have been a crucial component in phylogenetic analyses, they rarely are the key consideration; and in cases in which they are, the study is isolated in a single clade of organisms.

This report details a broad review of current knowledge of vertebrate teeth from to Mammalia, specifically focusing on their location, replacement, and attachment. Furthermore, this report also includes an analysis of a character map made from the current information on the teeth characteristics listed and on established vertebrate phylogenies. It is the hope of this author that by exploring the evolution of vertebrate teeth, a greater understanding of how teeth came to be, what they can become, and what could be done for man’s may be garnered by those who wish to know.

4

Introduction

Vertebrate evolution has been and continues to be researched and expanded upon for the past few decades. With advancements in multiple scientific fields such as , genetics, comparative anatomy, statistics, and phylogenetics, the world of vertebrate evolution—and evolution, in general—is rapidly expanding as researchers of those many fields integrate and assimilate their data to discover more wonders of this planet. Even in recent decades, many have explored vertebrate evolution in various ways. Blomme et al. (2006) traced evolution over the course of 600 million years, analyzing ancestral genes and the importance of gene duplication in the rise of complex ; Mindell and Honeycutt (1990) studied the exceptional possibility of utilizing ribosomal RNA to create a vertebrate phylogeny; and Romer

(1967) summarized the near entirety of vertebrate origins and speciation.

Though broad vertebrate evolution has been extensively researched, teeth (and specifically phylogenetics of teeth) have not been as explored. Teeth and their many variations have been a crucial element in paleontological and biological research in vertebrates for many years (Owen 1840). Current research on teeth is still voluminous, but the vast majority tends to focus on singular characteristics such as tooth location, attachment, replacement, genetics, and shape. For example, Davit-Beal et al. (2009) researched teeth loss in by reviewing current knowledge on teeth genetics and the record Smith and Johanson (2003) both studied the presence of teeth in derived placoderms known as athrodires through comparative anatomy, but they solely focused on athrodires, only referencing other vertebrates rather than giving them full analyses as well. Secondary sources such as Berkovitz and Shellis’s The Teeth of Non-Mammalian Vertebrates (2017) appear to be the few that analyze physical characteristics of teeth across all vertebrates as well as more microbiological ones. Lee 5

Phylogenetic analysis on teeth are of similar flavor to the more anatomical research.

Smith and Coates (1998) investigated the origins of teeth in vertebrates by analyzing denticles and odontodes, and their phylogenetic analyses revealed that teeth may have been formed deep within vertebrate diversification as an intricate modification to non-growing odontodes.

However, though they give suggestions and developmental models on how this new fact translates to the evolutionary biology of remaining vertebrates, they ultimately leave the task of further exploration to others. Fink (1981) also conducted phylogenetic research on vertebrates, but he focused specifically on tooth attachment in Actinopterygii. Similar treatment was given to hominids (Read 1975) and Amphibia (Parsons and Williams 1962), and any dental-based phylogenetic research focused on either one character, one clade, or one of each.

For this thesis, a comprehensive, -based review of all major classes and superclasses in vertebrates was conducted; and a character matrix and map of those classes were formed based on the information found in the review and on an appropriate phylogenetic tree.

Characters involved in the making of the matrix and map included location, degree of replacement, and mode of attachment since all of these have been thoroughly covered by others and are easily quantifiable compared to others. This research project ultimately attempts to ascertain how these three characters influenced speciation over evolutionary time. Through this thesis, the details of dental evolution may be elaborated, and further endeavors to analyze the whole of vertebrate dentition may be sparked.

6

Materials and Methods

With academic websites (Google Scholar, National Center for Biotechnology

Information, ScienceDirect, Web of Science, and JSTOR), primary and secondary sources pertaining to the location, presence of differentiation, degree of replacement, shape, and type of attachment of teeth for a vertebrate class or order were searched and collected. Phrases such as

“ teeth replacement” and “Amphibia teeth homodont/heterodont” were used in the websites to find said sources. After appropriate sources were obtained, any information that they had pertaining to the five aforementioned character categories were noted and arranged in a character matrix via Mesquite, a system used for evolutionary analysis (version 3.6). If a class, family, or order exhibited any presence of a character, it was marked on the matrix as a “1”—the absence of a character is marked as “0,” and unknown characters are marked as “?”. Once the character matrix was complete, Mesquite was used to determine synapomorphies and autapomorphies via parsimonious analysis and, ultimately, to map teeth characters onto a well- established vertebrate phylogeny.

All classes of extant vertebrates (Agnatha, Chondrichthyes, Osteichthyes, Amphibia,

Reptilia, Aves, and Mammalia) were studied and included in the character matrix and the phylogenetic tree, as well as the following subclasses, infraclasses, orders, and families:

• Orders Myxiniformes and Petromyzontiformes (Class Agnatha); • Subclasses Elasmobranchii and Holocephali (Class Chondrichthyes); • Subclass Chondrostei (Class Actinopterygii of Superclass Osteichthyes); • Order Polypteriformes (Class Actinopterygii of Superclass Osteichthyes); • Infraclasses Holostei and Teleostei (Class Actinopterygii of Superclass Osteichthyes); • Subclasses Actinistia and Dipnoi (Class Sarcopterygii of Superclass Osteichthyes); Lee 7

• Families Ambystomatidae, Amphiumidae, Cryptobranchidae, Hynobiidae, Proteidae, Salamandridae, Sirenidae, and Plethodontidae (Order Caudata of Class Amphibia); • Families Rhinatrematidae, , , Herpelidae, Caeciliidae, Typhlonectidae, Indotyphlidae, Siphonopedae, Dermophiidae, and Chikiidae (Order of Class Amphibia); • Families Pipidae, Bombinatoridae, Hemphractidae, Brachycephalidae, Hylidae, Ceratophrydae, Dendrobatidae, Ranidae, and Bufonidae(Order Anura of Class Amphibia); • Order Rhynchocephalia, Chelonii, and Crocodylia (Class Reptilia); • Families Chamaeleonidae, Dactyloidae, Agamidae, Iguanidae, Gekkonidae, Eublepharidae, Diplodactylidae, Pygopodidae, Teiidae, Lacertidae, Scincidae, Xantusiidae, Cordylidae, Amphisbaenia, Gerrhosauridae, Anguidae, Xenosauridae, Helodermatidae, Varanidae, Leptotyphlopidae, Typhlopidae, Pythonidae, Boidae, Lamprophiidae, Colubridae, Elapidae, and Viperidae (Order Squamata of Class Reptilia); • Orders Rodentia, Chiroptera, Soricomorpha, Primates, Carnivora, Artiodactyla, Diprotodontia, , Didelphimorphia, , Dasyuromorphia, Afrosoricida, Erinaceomorpha, Cingulata, Peramelemorphia, Scandentia, Perissodactyla, Macroscelidea, Pilosa, Monotremata, and Proboscidea (Class Mammalia); and • Orders Ornithischia and Saurischia (Class Dinosauria).

Regarding the teeth characters, “location” simply refers to where teeth are found in the clade. Focus was given on the marginal bones (, , dentary, coronoid, and splenial), the palatal bone, the pterygoid, the vomer, the , and the while noting any remarkable locations that could not be categorized by the aforementioned ones.

“Replacement” refers to how often organisms within a clade replaces its teeth. “Monophyodont” indicates that they never replace their teeth throughout their lifetime; “diphyodont” indicates one replacement cycle; “” indicates more than one cycle; and “elodont” indicates teeth 8 that continuously grow throughout their lives. Finally, “attachment” refers to how the teeth are attached to the bone, if they are attached to bone at all. “Thecodont” attachment means that the base of the tooth is embedded within a socket of bone. In contrast, “acrodont” attachment involves teeth that are attached to the alveolar ridge of the bones without sockets; and

“pleurodont” dentition involves teeth that are fused to the side of the jaw bones’ alveolar ridges.

The “ankylosis” category is designated to those clades that had no specific information regarding the type of ankylosis (thecodont, acrodont, or pleurodont). “Pedicellate” dentition is a special category for and —it entails teeth that have the typical and base, but they are separated by a layer of dentine. These teeth can also be ankylosed (Berkovitz and Shellis

2017). Finally, “connective” attachment refers to teeth that are not fused to bone but are instead connected to the via connective tissues like .

Lee 9

Results

Note: The character maps generated by the Mesquite program are found in Appendix 1, and their corresponding supplemental tables are in Appendix 2.

Location

Premaxillary and maxillary teeth are present in the vast majority of vertebrate classes and have multiple cases of synapomorphy in the phylogenetic trees. Synapomorphic gains in premaxillary teeth separated Chondrichthyes from Agnatha, Holostei and Teleostei from

Chondrostei, Tetrapoda (Amphibia, Reptilia, Aves, and Mammalia) from Dipnoi, most of

Caudata from Sirenidae, Dendrobatidae from Bufonidae, Pythonidae and Colubridae from many families, Dinosauria and Crocodylia from Aves and Chelonii, the rest of Mammalia from

Monotremata, and Cingulata from Pilosa (see Figure 1). Synapomorphies in maxillary teeth led to very similar outcomes with premaxillary teeth, but with a few more divisions. Gains in maxillary teeth divided Tetrapoda from both Actinistia and Dipnoi, Salamandridae and

Ambystomatidae from Proteidae, and Typhylopidae from Leptotyphlopidae; however, synapomorphic acquisition of maxillary teeth in Squamata separated the entire order from Aves

(Figure 2).

Synapomorphies involving dentary teeth led to the same outcomes as premaxillary teeth from Agnatha to Osteichthyes and within Gymnophiona of Amphibia, and as maxillary teeth within Squamata and Mammalia; however, a synapomorphic loss of these teeth divided Anurans from the rest of Amphibia and Typhlopidae from Leptotyphlopidae (Figure 3).

Unlike premaxillary, maxillary, and dentary teeth, coronoid and splenial teeth are much sparser in vertebrate evolution. Notable divisions caused by the presence of coronoid teeth include Polypteriformes from other Actinopterygii, Actinistia from Dipnoi and Tetrapoda, 10

Gymnophi ona from Anura and Caudata, and Amphibia from Reptilia and Mammalia. Proteidae,

Salamandridae, and Ambystomatidae of Caudata were also separated from Amphiumidae and

Plethodontidae as a result of coronoid dentition (Figure 4). Splenial dentition has even fewer synapomorphic influences—the most significant one divided Rhinatrematidae from the rest of

Amphibia (Figure 5).

Palatal teeth are non-existent in Agnatha and Chondrichthyes but are present in

Osteichthyes and beyond. Another significant synapomorphic division occured between

Amphibia (which retained palatal teeth) and Reptilia and Mammalia (which lost palatal teeth).

Rhynchocephalia’s acquisition of palatal teeth also separated them from the rest of Reptilia, and the more derived of Serpentes from Leptotyphlopidae and Typhlopidae (Figure 6).

Smaller divisions of families also occurred in Caudata and Anura.

A critical synapomorphy involving pterygoidal teeth can be found in Osteichthyes— divergence from Chondrichthyes to Osteichthyes was in part by development of those teeth. The second crucial synapomorphy involved the loss of pterygoidal teeth from Sarcopterygii

(Osteichthyes) to Tetrapoda. Furthermore, like the palatal teeth, the diverged snakes were separated from Leptotyphlopidae and Typhlopidae due to synapomorphic gains in pterygoidal teeth; and smaller divisions can be found in caudates and other squamates (Figure 7).

Similar to palatal and pterygoidal teeth, synapomorphic emergence of vomerine teeth separates Osteichthyes from Agnatha and Chondrichthyes. Two more major divisions are present as well—Reptilia and Mammalia had a synapomorphic loss in vomerine teeth, splitting them from Amphibia; and Amphibia also acquired a loss, separating them from Caudata (Figure 8).

Other synapomorphies are also present, but they involve individual, heavily diverged clades. Lee 11

Tongue teeth have only two synapomorphies total—one is found at the split between

Agnatha and Chondrichthyes, in which Agnatha bear the teeth; and the other is located at the

Holostei-Teleostei junction, in which Teleostei have them (Figure 9).

Pharyngeal teeth have one more synapomorphy compared to tongue teeth. First,

Petromyzontiformes acquired , differentiating it from the sister Chondrichthyes.

Next, Teleostei gained some pharyngeal teeth, dividing itself from other Actinopterygii. Finally,

Dipnoi gained pharyngeal teeth, splitting Osteichthyes from Tetrapoda (Figure 10).

Teeth in positions that could not be categorized by the above were found only in

Agnatha. Therefore, a synapomorphic loss of these kinds of teeth separated the other vertebrate classes from Agnatha (Figure 11).

Replacement

Monophyodonty emerged only in singular clades: Rhynchocephalia, Chamaeleonidae,

Diprotodontia, Cetacea, and Didelphimorphia. Although Chamaeleonidae and Cetacea are heavily diverged clades, the others relatively are not. Therefore, the emergence of monophyodonty split Rhynchocephalia from the other squamates and Didelphimorphia and

Diprotodontia from the more derived (Figure 12).

Diphyodonty exclusively arose in Mammalia as a synapomorphic gain separating them from Monotremata. Loss of diphyodonty appears in the more diverged clades of Pilosa,

Proboscidea, Cetacea, Lagomorpha, and Rodentia (Figure 13).

The synapomorphies of polyphyodonty, like the other replacement modes, are few; but they are involved in major evolutionary divisions. Loss of polyphyodonty distinguishes 12

Holocephali from Elasmobranchii; Dipnoi from Tetrapoda; and Mammalia from Reptilia (Figure

14).

Elodonty was phylogenetically involved in very few clades. Synapomorphic gains of this mode of replacement split Holocephali from Elasmobranchii and Dipnoi from Tetrapoda. Any other instances of elodonty are found in diverged clades of Mammalia (Pilosa, Rodentia, and

Lagomorpha) (Figure 15).

Attachment

Thecodont attachment primarily arose in Mammalia, signified by a synapomorphic gain dividing derived mammals from Monotremata. However, Crocodylia and Dinosauria also received a thecodont synapomorphy (which separated them from Squamata), and Teleostei gained thecodonty as well, separating them from Holostei (Figure 16).

A significant divergence involved a synapomorphic gain of acrodont attachment—

Amphibia split from Reptilia and Mammalia by the emergence of this form of attachment. Other significant divisions include a synapomorphic loss in Anura, Hynobiidae, and Cryptobranchidae

(which split them from Gymnophiona and more derived caudates); a synapomorphic gain in

Rhynchocephalia (which separated them from the rest of Reptilia); and a gain in the more speciated clades of Serpentes (which split them from Leptotyphlopidae and Typhlopidae). Small speciation events involving acrodonty can also be found in Squamata and Caudata (Figure 17).

Like acrodonty, pleurodonty also had a synapomorphic division between Amphibia and

Reptilia and Mammalia (Amphibia acquired pleurodonty). Another major division also occurred within Amphibia; specifically, Anura lost pleurodonty, dividing them from Caudata. Derived

Reptilia also split from Rhynchocephalia from a synapomorphic gain of acrodonty, and a Lee 13 synapomorphic loss divided derived clades in Serpentes from ancestral Leptotyphlopidae and

Typhlopidae. Smaller divisions are found in Caudata and Squamata (Figure 18).

Holostei and Teleostei acquired a synapomorphic gain of ankylosis that signified their derivation from Polypteriformes. The same occurred with Dipnoi and Tetrapoda. Smaller divisions happened in Caudata, Anura, and Squamata, with the most notable one being Pipidae’s emergence of ankylosis, separating the family from the rest of the anurans (Figure 19).

Pedicellate attachment initially exhibited “to-and-fro” behavior in Osteichthyes, first presenting as a synapomorphic gain in Polypteriformes, Teleostei, and Actinistia, but then becoming a loss for Dipnoi. However, Amphibia obtained pedicellate teeth, which distinguished them from Reptilia and Mammalia. Like ankylosis, smaller divisions were found in Squamata,

Caudata, and Anura, the most notable being Pipidae again (Figure 20).

Finally, connective attachment had only one critical synapomorphic event: the loss found in Osteichthyes. Agnatha and Chondrichthyes were the only ones with this mode of attachment

(Figure 21).

14

Discussion

Regarding locations of teeth, based on the information from the character maps, premaxillary, maxillary, and dentary teeth bore the most prominent role in the speciation of dentition in Vertebrata. Starting from Chonrichthyes, these three teeth locations pervaded the vast majority of clades and often correlated with major speciation events that separated even entire orders and classes of organisms. Such a surge of teeth in these locations correlates with the evolution of jawed vertebrates from Agnatha to Chondrichthyes, which is one of the more hypothesized theories regarding the origins of teeth (McCollum and Sharpe 2001). Smith (2003) also noted a similar fact, proposing that the developmental systems responsible for regulating pharyngeal teeth (or bone-related teeth) in jawless Agnatha also influenced the emergence of marginal teeth. Other marginal locations, on the other hand, were not as involved—as covered before, the clades that are influenced by coronoid and especially splenial teeth are few.

Patterning of these teeth in the character maps indicates that they were lost over evolutionary time, which coincides with a trend of increasing dental reduction in more diverged classes as their teeth transitioned from scattershot placements in the to locations more suited to their unique dietary needs (Huysseune and Sire 1998).

In fact, this reductive trend can be applied to the other non-marginal locations as well.

Both tongue and pharyngeal teeth had remarkably short evolutionary lifespans ending at

Osteichthyes. Palatal and pterygoidal teeth are present in Amphibia and Reptilia; however, palatal teeth mostly disappear in the more advanced Caudata and Anura, and both palatal and pterygoidal teeth are absent in Reptilia except for a few clades in Squamata, most notably the more derived Serpentes. Vomerine teeth are missing in Reptilia (except for Anguidae) but are present in Amphibia, but in Amphibia, almost all anuran clades lack these teeth. Considering that Lee 15 most of the amphibians come from Anura, losing vomerine teeth correlates with the increase in speciation events in Amphibia. Therefore, teeth locations are intrinsically tied to vertebrate evolution in that as more speciation events and divergences occurred, the more reduced the number of tooth locations became.

Teeth replacement is an extremely well-conserved characteristic in vertebrate evolution.

All classes, even one clade in Mammalia, exhibited some degree of polyphyodonty; and besides mammals, very few clades have different forms of replacement. Only two families in Reptilia

(Chamaeleonidae and Rhynchocephalia) and three in Mammalia (Diprotodontia, Cetacea, and

Didelphimorphia) lacked any replacement at all.

Based on the character maps, polyphyodonty generally behaves similarly to teeth locations—as vertebrates became more speciated, polyphyodonty reduced over time (albeit it was an extremely long time before a class of vertebrates adopted reduced polyphyodonty).

Furthermore, unlike diphyodonty and polyphyodonty, monophyodonty and elodonty (technically a special variant of monophyodonty) are scattered throughout the tree, present only in individual clades. This observation reinforces both the trend of reduced polyphyodonty and the conservation of teeth replacement—namely, instances of monophyodonty and elodonty are remarkably scarce (a sign of replacement conservation), but cases in which they do occur involve specific speciation of a few clades usually deep within the phylogeny (a sign of the reduced polyphyodonty trend).

These various degrees of replacement have been discussed, specifically regarding how these came to be. Osborn (1975) states that the difference in replacement rates between Reptilia and Mammalia were related to teeth efficiency. In other words, the need for greater replacement depends on whether or not the teeth are appropriate for the at that specific point in time. 16

If an animal undergoes rapid and constant change in size or morphology, or if the usage of teeth changes, polyphyodonty is necessary to accommodate these changes. Related to this is the research on dental lamina conducted by Tucker and Fraser (2014), in which they determined that the dental lamina (epithelial tissue responsible for redeveloping teeth) was a conserved trait in tooth regeneration, but the genes expressed are different in each class. Both of these observations support the trends observed in the character maps—more speciated and derived organisms with more complex lives and diets require teeth replacement, but simultaneously the teeth are more specialized not only to accommodate the lifestyles but to maximize efficiency and reduce the need for excessive replacement.

Finally, attachment does not appear to follow an overall linear trend like location and replacement do. Instead, this category of characters is perhaps the most “specialized” of the three because each of the attachment types seemingly belongs to specific classes or orders. Connective attachment is found only in Agnatha and Chondrichthyes; pedicellate attachment is mainly in

Actinopterygii and Amphibia; thecodont attachment is in Crocodylia, Dinosauria, and

Mammalia; acrodonty is primarily in Gymnophiona, Caudata, and Serpentes; and pleurodonty is mainly found in Gymnophiona, Caudates, and Squamata (excluding Serpentes).

However, determinations on which attachment types are more ancestral or derived can be made through the trees; and thus, an observation on how teeth attachment evolved can be gleaned. Based on the maps, the following attachment types are arranged from most ancestral to most derived: connective, pedicellate, ankylosis, pleurodont, acrodont, and thecodont. (The reason for acrodonty’s position over pleurodonty is that whereas pleurodonty influenced , acrodonty influenced snakes, which are more derived than lizards. The rest are relatively easy to trace.) This was to be expected since, like the vertebrates themselves, the modes of attachment as Lee 17 listed become more physiologically advanced. Therefore, according to the data and maps, there is a direct relationship between attachment complexity and organismal speciation.

Variance in attachment modes has been covered heavily by Gaengler (2000) and

Huysseune and Sire (1998). Gaengler (2000) dictates that significant speciation in vertebrates demanded improvements to tooth attachment modes as teeth became more varied and specialized. Moreover, he emphasizes the importance of the selection of socketed teeth by mammals, which deviated from the pleurodont Reptilia. Both of these statements correlate with what was seen in the character maps—overall, as more speciation events occurred, the types of attachment mode became more complex, starting from simple attachment via ligamentous tissue to socketed attachment with multiple connective tissues and fused bones. Huysseune and Sire

(1998) discuss the origins of the attachment bones, questioning whether or not they came from the teeth or from bony structures. They also mention that the attachment bones could have led to the development of the mammalian alveolar bone and, ultimately, thecodont attachment.

Although the character maps do not provide an answer to this question, they do depict the importance of the attachment bones as clades diverged. Without those bones, advanced attachment and teeth specialization may not have been possible.

Although the character maps produced in this research provided valuable information regarding location, replacement, and attachment, they are ultimately based on a relatively generalized phylogenetic tree. Individual genera, which could offer a more in-depth breakdown of the three characters, were not analyzed for this project; thus, the trees are susceptible to being too generalized. Furthermore, all information for the trees were based on metanalysis of multiple sources from other researchers. Therefore, as time progresses, new information can affect how these characters differentiate in Vertebrata. If possible, an expansion on this project involving 18 genera and new data as they come is recommended, as continued research on these characters, as shown in this project, can offer a considerable wealth of information regarding vertebrate teeth evolution. Finally, one last expansion can include other characteristics such as shape and differentiation so that the evolutionary story of teeth is more complete.

Lee 19

Conclusion

This project, a mapping of tooth locations, replacement, and attachment on a phylogeny of vertebrates, revealed remarkable trends of divergence and speciation in the three characters.

Both the number of locations in which teeth were present and the number of times that they were replaced generally declined over evolutionary time and as species diverged, whereas the mode of attachment gradually became more complex. Further and more in-depth research in the future years is encouraged for any follow-ups of this project, as it could provide a more complete history of dental evolution in vertebrates.

20

References

Ahlberg PE, Smith MM, Johanson Z. 2006. Developmental plasticity and disparity in early dipnoan (lungfish) . Evol Dev. 8:331-349.

Berkovitz BKB., Shellis RP. 2017. The Teeth of Non-Mammalian Vertebrates. London: Academic Press.

_____, _____. 2018. The Teeth of Mammalian Vertebrates. London: Academic Press.

_____, Sloan P. 1979. Attachment tissues of the teeth in Caiman sclerops (). J Zool Lond. 187:179-194.

Blomme T, Vandepoele K, De Bodt S, Simillion C, Maere S, Van de Peer Y. 2006. The gain and loss of genes during 600 million years of vertebrate evolution. Genome Biol. 7(5):R43.

Buchtova M, Zahradnicek O, Balkova S, Tucker AS. 2013. Odontogenesis in the Veiled Chameleon (Chamaeleo calyptratus). Arch Oral Biol. 58(2):118-133.

Carr SM. 2013. Traditional versus Phylogenetic classification of Vertebrata. http://www.mun.ca/biology/scarr/Vertebrate_phylogeny.html.

Cavin L, Valentin X, Garcia G. 2016. A new mawsoniid coelacanth (Actinistia) from the Upper Creatceous of Southern France. Cretac Res. 62:65-73.

Chiari Y, Cahais V, Galtier N, Delsuc F. 2012. Phylogenomic analyses support the position of turtles as the sister group of birds and (Archosauria). BMC Biol. 10(65):65.

Clemen G, Bartsch P, Wacker K. 1998. Dentition and dentigerous bones in juvenile adults of Polypterus senegalus (Cladistia, Actinopterygii). Ann Anat. 180:211-221.

Cooper JS, Poole DFG, Lawson R. 1970. The dentition of agamid lizards with special reference to tooth replacement. J Zool Lond. 182:85-98.

Davit-Beal T, Chisaka H, Delgado S, Sire J-Y. 2007. teeth: current knowledge, unanswered questions, and some directions for future research. Biol Rev. 92:49-81.

Degener LM. 1924. The development of the dentary teeth of Amia calva. J Morphol. 39:113-139.

Delgado S, Davit-Beal T, Sire J-Y. 2003. Dentition and Tooth Replacement Pattern in Chalcides (Squamata; Scincidae). J Morphol. 256(2):146-159. Lee 21

Enax J, Fabritius H-O, Rack A, Prymak O, Raabe D, Epple M. 2013. Characterization of teeth: Correlation of composition, microstructure, and hardness. J Struct Biol. 184:155- 163.

Fraser GJ, Berkovitz BK, Graham A, Smith MM. 2006. Gene deployment for tooth replacement in the rainbow trout (Oncorhynchus mykiss): a developmental model for evolution of the osteichthyan dentition. Evol Dev. 8:446-457.

Fink WL. 1981. Ontogeny and phylogeny of tooth attachment in actinopterygian . J Morphol 167(2):167-184.

Frazzetta TH. 1988. The mechanics of cutting and the form of teeth (Chondrichthyes, Elasmobranchii). Zoomorphology. 108:93-107.

Frost DR, Grant T, Faivovich J, Bain RH, Haas A, Haddad CFB, De Sa RO, Channing A, Wilkinson M, Donnellan SC, Raxworthy CJ, Campbell JA, Blotto BL, Moler P, Drewes RC, Nussbaum RA, Lynch JD, Green DM, Wheeler WC. 2006. The Amphibian Tree of Life. Bull Am Mus Nat Hist. 2006(297):1-291.

Gaengler P. 2000. Evolution of tooth attachment in lower vertebrates to tetrapods. In: Teaford MF, Smith MM, Ferguson MWJ, editors. Development, Function and Evolution of Teeth. New York, NY: Cambridge University Press.

Grady JE. 1970. Tooth Development in Latimeria chalumnae. J Morphol. 132:377-388.

Hardistry MW. 1979. Biology of the Cyclostomes. New York (NY): Springer.

Helfman G, Collette BB, Facey DE, Bowen BW. 2009. The Diversity of Fishes: Biology, Evolution, and Ecology. West Sussex, UK: Wiley-Blackwell.

Huysseune A, Sire J-Y. 1998. Evolution of patterns and processes in teeth and tooth-related tissues in non-mammalian vertebrates. Eur J Oral Sci. 106:437-481.

Huysseune A, Witten PE. 2006. Developmental Mechanisms Underlying Tooth Patterning in Continuously Replacing Osteichthyan Dentitions. J Exp Zool. 306B:204-215.

Jenkins KM, Jones MEH, Zikmund T, Boyde A, Daza JD. 2017. A Review of Tooth Implantation Among Rhynchocephalians (Lepidosauria). J Herpetol. 51(3):300-306.

22

Kerr T. 1960. Development and Structure of Some Actinopterygian and Urodele Teeth. Proc Zool Soc Lond. 133:401-422.

LeBlanc ARH, Brink KS, Cullen TM, and Reisz RR. 2017. Evolutionary implications of tooth attachment versus tooth implantation: A case study using , crocodilian, and teeth. J Vertebr Paleontol. e1354006-1-19.

Louchart A, Viriot L. 2011. From snout to : the loss of teeth in birds. Trends Ecol Evol. 26:663-673.

Mahler DL, Kearney M. 2006. The Palatal Dentition in Squamate : Morphology, Development, Attachment, and Replacement. . 108:1-61.

Martin KJ, Rasch LJ, Cooper RL, Metscher BD, Johanson Z, Fraser GJ. 2016. Sox2+ progenitors in link taste development with the evolution of regenerative teeth from denticles. PNAS 113(51):14769-14774.

Mateo JA, Lopez-Jurado LF. 1997. Dental Ontogeny in Lacerta lepida (Sauria, Lacertidae) and Its Relationship to Diet. Copeia. 1997(2):461-463.

McCollum M, Sharpe PT. Evolution and development of teeth. J Anat. 199:153-159.

Meunier FJ, Mondejar-Fernandez J, Goussard F, Clement G, and Herbin M. 2015. Presence of plicidentine in the oral teeth of the coelacanth Latimeria chalumnae Smith 1939. (Sarcopterygii; Actinistia). J Struct Biol. 190:31-37.

Miller WA, Radnor CJP. 1973. Tooth Replacement in the Bowfin (Amia calva - Holostei). J Morphol. 140(4):381-395.

Mindell DP, Honeycutt RL. 1990. Ribosomal RNA in Vertebrates: Evolution and Phylogenetic Applications. Annu Rev Ecol Syst. 21:541-566.

Moldowan PD, Brooks RJ, Litzgus JD. 2016. Turtles with "teeth": beak morphology of Testudines with a focus on the tomiodonts of Painted Turtles (Chrysemys spp.). Zoomorphology. 135:121-135.

Nance HA. 2007. Cranial osteology of the African gerrhosaurid Angolosaurus skoogi (Squamata; Gerrhosauridae). Afr J Herpetol. 56(1):39-75.

Lee 23

Nikitina NG, Ananjeva NB. 2009. Characteristics of Dentition in Gekkonid Lizards of the Genus Teratoscincus and Other Gekkota (Sauria, Reptilia). Biol Bulletin. 36(2):193-198.

Osborn JW. 1975. Tooth Replacement: Efficiency, Patterns and Evolution. Evolution 29(1):180- 186.

Owen R. 1840. Odontography; or, A treatise on the comparative anatomy of the teeth; their physiological relations, mode of development, and microscopic structure of the vertebrate . H. Bailliere, London.

Parsons TS, Williams EE. 1962. The Teeth of Amphibia and Their Relation to Amphibian Phylogeny. J Morphol 110(3):375-389.

Presch W. 1974. A Survey of the Dentition of the Macroteiid Lizards (Teiidae: Lacertilia). Herpetologica. 30(4):344-349.

Qu Q, Sanchez S, Blom H, Tafforeau P, Ahlberg PE. 2013. Scales and Tooth Whorls of Ancient Fishes Challenge Distinction between External and Oral 'Teeth.' PLOS ONE 8.

Read DW. 1975. Hominid teeth and their relationship to hominid phylogeny. Am J Phys Anthropol. 42(1):105-125.

Richman JM, Handrigan GR. 2011. Reptilian Tooth Development. genesis. 49(4):247-260.

Rieppel O. 1980. The Phylogeny of Anguinomorph Lizards. Basel (Switzerland): Birkhäuser Basel.

Rocek Z. 1980. The Dentition of the European Glass Ophisaurus apodus (Pallas, 1775) (Reptilia, Sauria: Anguidae), with Notes on the Pattern of Tooth Replacement. Amphibia- Reptilia. 1(1):19-27.

Romer AS. 1956. Osteology of the Reptiles. Chicago (IL): University of Chicago Press.

_____. 1967. Major Steps in Vertebrate Evolution. Science 158(3809):1629-1637.

_____, Parsons TS. 1986. The Vertebrate Body. 6th ed. New York (NY): Saunders College Publishing.

Sander PM. 1997. Teeth and Jaws. In: Currie PJ, Padian K, editors. The Encyclopedia of . Cambridge, MA: Academic Press. 24

Schultze H-P. 1986. Dipnoans as Sarcopterygians. J Morphol. 190:39-74.

Schwenk K, editor. 2000. Feeding: Form, Function, and Evolution in Vertebrates. Cambridge, MA: Academic Press.

Serano PC. 1997. The Origin and Evolution of Dinosaurs. Annu Rev Earth Planet Sci. 25:435- 489.

Smirnov SV, Vasil'eva AB. 1995. Anuran Dentition: Development and Evolution. Russ J Herpetol. 2:120-128.

Smith JLB. 1939. A SURVIVING FISH OF THE ORDER ACTINISTIA. Transactions of the Royal Society of South Africa 27:47-50.

Smith MM. 2003. Verebrate dentitions at the origins of jaws: when and how pattern evolved. Evol Dev. 5(4):394-413.

_____, Coates MI. 1998. Evolutionary origins of the vertebrate dentition: phylogenetic patterns and developmental evolution. Eur J Oral Sci. 106(suppl 1):482-500.

_____, Johanson Z. 2003. Separate Evolutionary Origins of Teeth in Fossil Jawed Vertebrates. Science 299(5610):1235-1236.

Tucker AS, Fraser GJ. 2014. Evolution and developmental diversity of tooth regeneration. Semin Cell Dev Biol. 25-26:71-80.

Underwood CJ, Johanson Z, Welten M, Metscher B, Rasch LJ, Fraser GJ, Smith MM. 2015. Development and Evolution of Dentition Pattern and Tooth Order in the Skates and Rays (Batoidea; Chondricthyes). PLOS ONE.

Ungar PS. 2010. Mammal Teeth: Origin, Evolution, and Diversity. Baltimore, MD: Johns Hopkins University Press.

Van der Heyden C, Huysseune A, Sire J-Y. 2000. Development and fine structure of pharyngeal replacement teeth in juvenile zebrafish (Danio rerio) (Teleostei, Cyprinidae). Cell Tissue Res. 302:205-219.

Wake MH. 1976. The Development and Replacement of Teeth in Viviparous . J Morphol. 148:33-64. Lee 25

Whitlock JA, and Richman JM. 2013. Biology of tooth replacement in amniotes. Int J Oral Sci. 5(2):66-70.

Wiens JJ, Hutter CR, Mulcahy DG, Noonan BP, Townsend TM, Sites JW, Reeder TW. 2012. Resolving the phylogeny of lizards and snakes (Squamata) with extensive sampling of genes and species. Biol Lett. 8(6):1043-1046.

Zaher H, Rieppel O. 1999. Tooth Implantation and Replacement in Squamates, with Special Reference to Mosasaur Lizards and Snakes. American Museum Novitates. 3271.

26

Appendix 1: Character Maps

Note: The phylogenetic tree used to generate the character maps was constructed based on data from Carr (2013), Chiari et al. (2012), Frost et al. (2006), and Wiens et al. (2012).

Figure 1. Premaxillary teeth in vertebrates.

Figure 2. Maxillary teeth in vertebrates. Lee 27

Figure 3. Dentary teeth in vertebrates.

Figure 4. Coronoid teeth in vertebrates.

28

Figure 5. Splenial teeth in vertebrates.

Figure 6. Palatal teeth in vertebrates.

Lee 29

Figure 7. Pterygoidal teeth in vertebrates.

Figure 8. Vomerine teeth in vertebrates.

30

Figure 9. Tongue teeth in vertebrates.

Figure 10. Pharyngeal teeth in vertebrates.

Lee 31

Figure 11. Other teeth locations in vertebrates.

32

Figure 12. Monophyodonty in vertebrates.

Figure 13. Diphyodonty in vertebrates.

Lee 33

Figure 14. Polyphyodonty in vertebrates.

Figure 15. Elodonty in vertebrates.

34

Figure 16. Thecodont attachment in vertebrates.

Figure 17. Acrodont attachment in vertebrates.

Lee 35

Figure 18. Pleurodont attachment in vertebrates.

Figure 19. Ankylosis in vertebrates.

36

Figure 20. Pedicellate attachment in vertebrates.

Figure 21. Connective attachment in vertebrates.

Lee 37

Appendix 2: Supplemental Tables

Supplemental Table 1.1. Tooth Locations in Agnatha, Chondrichthyes, Osteichthyes, and Amphibia (Caudata and Gymnophiona). “1” = Yes, “0” = No, “?” = Unknown, “-” = Not Applicable. Pre. = premaxilla, Max. = maxilla, Den. = dentary, Cor. = coronoid, Spl. = splenial, Pal. = , Pt. = pterygoid, Vom. = vomer, Ton. = tongue, Ph. = pharyngeal.

Pre. Max. Den. Cor. Spl. Pal. Pt. Vom. Ton. Ph. Other References AGNATHA Myxiniformes 0 0 0 0 0 0 0 0 1 0 1 Berkovitz and Shellis Petromyzontiformes 0 0 0 0 0 0 0 0 1 1 1 2017, Hardistry 1979. CHONDRICHTHYES Elasmobranchii 1 1 1 0 0 0 0 0 0 0 0 Berkovitz and Shellis Holocephali 1 1 1 0 0 0 0 0 0 0 0 2017, Frazzetta 1988, Huysseune and Sire 1998, Qu et al. 2013, Romer and Parsons 1986, Underwood et al. 2015. OSTEICHTHYES ACTINOPTERYGII Chondrostei ------Berkovitz and Shellis Polypteriformes 1 1 1 1 0 1 1 1 0 0 0 2017, Clemen et al. Holostei 1 1 1 0 0 1 1 1 0 0 0 1998, Degener 1924, Van der Heyden et al. 1 1 1 0 0 1 1 1 1 1 0 Teleostei 2000. SARCOPTERYGII Actinistia 1 0 1 1 0 1 1 1 0 0 0 Ahlberg et al. 2006, Dipnoi 0 0 0 0 0 1 1 1 0 1 0 Berkovitz and Shellis 2017, Cavin et al. 2016, Huysseune and Sire 1998, Meunier et al. 2015, Schultze 1986, Smith 1939. AMPHIBIA CAUDATA Ambystomatidae 1 1 1 1 1 1 0 1 0 0 0 Amphiumidae 1 1 1 0 0 0 0 1 0 0 0 Berkovitz and Shellis Cryptobranchidae 1 1 1 0 0 0 0 1 0 0 0 2017, Davit-Beal et al. Hynobiidae 1 1 1 0 0 0 0 1 0 0 0 2007, Huysseune and Proteidae 1 0 1 1 0 1 1 1 0 0 0 Sire 1998, Tucker and Salamandridae 1 1 1 1 1 0 0 1 0 0 0 Fraser 2014, Wake Sirenidae 0 0 0 0 1 1 0 1 0 0 0 1976. Plethodontidae 1 1 1 0 0 0 0 1 0 0 0 GYMNOPHIONA Rhinatrematidae 1 1 1 1 1 1 0 1 0 0 0 Ichthyophiidae 1 1 1 1 0 1 0 1 0 0 0 Scolecomorphidae 1 1 1 1 0 1 0 1 0 0 0 Herpelidae 1 1 1 1 0 1 0 1 0 0 0 Caeciliidae 1 1 1 1 1 1 0 1 0 0 0 Berkovitz and Shellis 2017, Davit-Beal et al. Typhylonectidae 1 1 1 1 1 1 0 1 0 0 0 2007. Indotyphlidae 1 1 1 1 1 1 0 1 0 0 0 Siphonopedae 1 1 1 1 0 1 0 1 0 0 0 Dermophiidae 1 1 1 1 0 1 0 1 0 0 0 Chikiidae 1 1 1 1 0 1 0 1 0 0 0

38

Supplemental Table 1.2. Tooth Locations in Amphibia (Anura), Reptilia, and Aves. Pre. Max. Den. Cor. Spl. Pal. Pt. Vom. Ton. Ph. Other References AMPHIBIA ANURA Pipidae 1 1 0 0 0 0 0 0 0 0 0 Bombinatoridae 1 1 0 0 0 0 0 0 0 0 0 Hemiphractidae 1 1 1 0 0 0 0 1 0 0 0 Brachycephalidae 1 1 0 0 0 0 0 0 0 0 0 Berkovitz and Shellis Hylidae 1 1 0 0 0 0 0 0 0 0 0 2017, Davit-Beal et al. Ceratophrydae 1 1 0 0 0 1 0 0 0 0 0 2007. Dendrobatidae 1 1 0 0 0 0 0 0 0 0 0 Ranidae 1 1 0 0 0 0 0 1 0 0 0 Bufonidae ------REPTILIA RHYNCHOCEPHALIA 1 1 1 0 0 1 0 0 0 0 0 Berkovitz and Shellis 2017. CHELONII ------Moldowan et al. 2016. CROCODYLIA 1 1 1 0 0 0 0 0 0 0 0 Berkovitz and Shellis 2017, Tucker and Fraser 2014. SQUAMATA Chamaeleonidae 1 1 1 0 0 0 0 0 0 0 0 Dactyloidae 1 1 1 0 0 0 0 0 0 0 0 Agamidae 1 1 1 0 0 0 0 0 0 0 0 Iguanidae 1 1 1 0 0 1 1 0 0 0 0 Gekkonidae 1 1 1 0 0 0 0 0 0 0 0 Eublepharidae 1 1 1 0 0 0 0 0 0 0 0 Diplodactylidae 1 1 1 0 0 0 0 0 0 0 0 Pygopodidae 1 1 1 0 0 0 0 0 0 0 0 Teiidae 1 1 1 0 0 0 1 0 0 0 0 Lacertidae 1 1 1 0 0 0 1 0 0 0 0 Scincidae 1 1 1 0 0 0 1 0 0 0 0 Berkovitz and Shellis 2017, Delgado et al. Xantusiidae 1 1 1 0 0 0 0 0 0 0 0 2003, Mahler and Cordylidae 1 1 1 0 0 0 0 0 0 0 0 Kearney 2006, Mateo Gerrhosauridae 1 1 1 0 0 0 1 0 0 0 0 and Lopez-Jurado 1997, Anguidae 1 1 1 0 0 1 1 1 0 0 0 Nance 2007, Presch Xenosauridae 1 1 1 0 0 0 0 0 0 0 0 1974, Rieppel 1980, Helodermatidae 1 1 1 0 0 1 1 0 0 0 0 Rocek 1980, Romer 1956. Varanidae 1 1 1 0 0 0 0 0 0 0 0 Amphisbaenia 1 1 1 0 0 0 0 0 0 0 0 Leptotyphlopidae 0 0 1 0 0 0 0 0 0 0 0 Typhlopidae 0 1 0 0 0 0 0 0 0 0 0 Pythonidae 1 1 1 0 0 1 1 0 0 0 0 Boidae 0 1 1 0 0 1 1 0 0 0 0 Lamprophiidae 0 1 1 0 0 1 1 0 0 0 0 Colubridae 1 1 1 0 0 1 1 0 0 0 0 Elapidae 0 1 1 0 0 1 1 0 0 0 0 Viperidae 0 1 1 0 0 1 1 0 0 0 0 AVES ------Louchart and Viriot 2011.

Lee 39

Supplemental Table 1.3. Tooth Locations in Mammalia and Dinosauria.

Pre. Max. Den. Cor. Spl. Pal. Pt. Vom. Ton. Ph. Other References MAMMALIA Rodentia 1 1 1 0 0 0 0 0 0 0 0 Chiroptera 1 1 1 0 0 0 0 0 0 0 0 Soricidae 1 1 1 0 0 0 0 0 0 0 0 Primates 1 1 1 0 0 0 0 0 0 0 0 Carnivora 1 1 1 0 0 0 0 0 0 0 0 Artiodactyla 1 1 1 0 0 0 0 0 0 0 0 Dirptodontia 1 1 1 0 0 0 0 0 0 0 0 Lagomorpha 1 1 1 0 0 0 0 0 0 0 0 Didelphimorphia 1 1 1 0 0 0 0 0 0 0 0 Cetacea 1 1 1 0 0 0 0 0 0 0 0 Berkovitz and Shellis Dasyuromorphia 1 1 1 0 0 0 0 0 0 0 0 2018, Ungar 2010. Afrosoricida 1 1 1 0 0 0 0 0 0 0 0 Erinaceomorpha 1 1 1 0 0 0 0 0 0 0 0 Cingulata 0 1 1 0 0 0 0 0 0 0 0 Peramelemorphia 1 1 1 0 0 0 0 0 0 0 0 Scandentia 1 1 1 0 0 0 0 0 0 0 0 Perissodactyla 1 1 1 0 0 0 0 0 0 0 0 Macroscelidea 1 1 1 0 0 0 0 0 0 0 0 Pilosa 1 1 1 0 0 0 0 0 0 0 0 Monotremata ------Proboscidea 1 1 1 0 0 0 0 0 0 0 0 DINOSAURIA Ornithischia 1 1 1 0 0 0 0 0 0 0 0 Sander 1997, Serano Saurischia 1 1 1 0 0 0 0 0 0 0 0 1997.

40

Supplemental Table 2.1. Tooth Replacement and Attachment in Agnatha, Chondrichthyes, Osteichthyes, and Amphibia (Caudata and Gymnophiona). “1” = Yes, “0” = No, “?” =

Unknown, “-” = Not Applicable. Mp. = monophyodont, Dp. = diphyodont, Pp. = polyphyodont, Elo. = elodont, Th. = thecodont, Acr. = acrodont, Pl. = pleurodont, Ank. = ankylosis, Ped. = pedicellate, Con. = connective.

REPLACEMENT ATTACHMENT References Mp. Dp. Pp. Elo. Th. Acr. Pl. Ank. Ped. Con. AGNATHA Myxiniformes 0 0 1 0 0 0 0 0 0 1 Berkovitz and Shellis Petromyzontiformes 0 0 1 0 0 0 0 0 0 1 2017, Hardistry 1979. CHONDRICHTHYES Elasmobranchii 0 0 1 0 0 0 0 0 0 1 Berkovitz and Shellis Holocephali 0 0 0 1 0 0 0 0 0 1 2017, Helfman et al. 2009, Martin et al. 2016, Romer and Parsons 1986, Underwood et al. 2015. OSTEICHTHYES ACTINOPTERYGII Chondrostei ------Berkovitz and Shellis Polypteriformes 0 0 1 0 0 0 0 0 1 0 2017, Clemen et al. Holostei 0 0 1 0 0 0 0 1 0 0 1998, Fink 1981, Fraser et al. 2006, Kerr 1960, 0 0 1 0 1 0 0 1 1 0 Teleostei Miller and Radnor 1973, Van der Heyden et al. 2000. SARCOPTERYGII Actinistia 0 0 1 0 0 0 0 0 1 0 Berkovitz and Shellis Dipnoi 0 0 0 1 0 0 0 1 0 0 2017, Cavin et al. 2016, Grady 1970, Huysseune and Sire 1998, Meunier et al. 2015. AMPHIBIA CAUDATA Ambystomatidae 0 0 1 0 0 1 1 0 1 0 Amphiumidae 0 0 1 0 0 1 1 ? ? ? Cryptobranchidae 0 0 1 0 0 0 1 0 1 0 Hynobiidae 0 0 1 0 0 0 1 0 1 0 Berkovitz and Shellis 2017, Davit-Beal et al. Proteidae 0 0 1 0 0 1 1 0 1 0 2007. Salamandridae 0 0 1 0 0 0 1 0 1 0 Sirenidae 0 0 1 0 0 1 0 1 0 0 Plethodontidae 0 0 1 0 0 0 1 0 1 0 GYMNOPHIONA Rhinatrematidae 0 0 1 0 0 1 1 0 1 0 Ichthyophiidae 0 0 1 0 0 1 1 0 1 0 Scolecomorphidae 0 0 1 0 0 1 1 0 1 0 Herpelidae 0 0 1 0 0 1 1 0 1 0 Caeciliidae 0 0 1 0 0 1 1 0 1 0 Berkovitz and Shellis 2017, Davit-Beal et al. Typhylonectidae 0 0 1 0 0 1 1 0 1 0 2007, Wake et al. 1976. Indotyphlidae 0 0 1 0 0 1 1 0 1 0 Siphonopedae 0 0 1 0 0 1 1 0 1 0 Dermophiidae 0 0 1 0 0 1 1 0 1 0 Chikiidae 0 0 1 0 0 1 1 0 1 0

Lee 41

Supplemental Table 2.2. Tooth Replacement and Attachment in Amphibia (Anura), Reptilia and Aves. REPLACEMENT ATTACHMENT References Mp. Dp. Pp. Elo. Th. Acr. Pl. Ank. Ped. Con. AMPHIBIA ANURA Pipidae 0 0 1 0 0 0 0 1 0 0 Bombinatoridae 0 0 1 0 0 0 0 0 1 0 Hemiphractidae 0 0 1 0 ? ? ? ? ? ? Brachycephalidae 0 0 1 0 ? ? ? ? ? ? Berkovitz and Shellis 2017, Davit-Beal et al. Hylidae 0 0 1 0 ? ? ? ? ? ? 2007, Smirnov and Ceratophrydae 0 0 1 0 0 0 0 1 0 0 Vasil'eva 1995. Dendrobatidae 0 0 1 0 0 0 0 0 1 0 Ranidae 0 0 1 0 0 0 0 0 1 0 Bufonidae ------REPTILIA RHYNCHOCEPHALIA 1 0 1 0 0 1 0 0 0 0 Berkovitz and Shellis 2017, Jenkins et al. 2017. CHELONII ------Moldowan et al. 2016. CROCODYLIA 0 0 1 0 1 0 0 0 0 0 Berkovitz and Shellis 2017, Berkovitz and Sloan 1979, Enax et al. 2013, Huysseune and Sire 1998, Huysseune and Witten 2006, Richman and Handrigan 2011. SQUAMATA Chamaeleonidae 1 0 0 0 0 1 0 0 0 0 Dactyloidae 0 0 1 0 0 0 1 0 0 0 Agamidae 0 0 1 0 0 1 1 0 0 0 Iguanidae 0 0 1 0 0 0 1 0 0 0 Gekkonidae 0 0 1 0 0 0 1 0 0 0 Eublepharidae 0 0 1 0 0 0 1 0 0 0 Diplodactylidae 0 0 1 0 0 0 1 0 0 0 Pygopodidae 0 0 1 0 0 0 0 1 1 0 Berkovitz and Shellis Teiidae 0 0 1 0 0 0 1 0 0 0 2017, Buchtova et al. Lacertidae 0 0 1 0 0 0 1 1 0 0 2013, Cooper et al. 1970, Scincidae 0 0 1 0 0 0 1 1 0 0 Delgado et al. 2003, Xantusiidae 0 0 1 0 0 0 1 0 0 0 Mahler and Kearney Cordylidae 0 0 1 0 0 0 1 0 0 0 2006, Mateo and Lopez- Jurado 1997, Nance 2007, Gerrhosauridae 0 0 1 0 0 0 1 0 0 0 Nikitina and Ananjeva Anguidae 0 0 1 0 0 0 1 0 0 0 2009, Presch 1974, Xenosauridae ? ? ? ? ? ? ? ? ? ? Rieppel 1980, Rocek Helodermatidae 0 0 1 0 0 0 1 0 0 0 1980, Romer 1956, Varanidae 0 0 1 0 0 0 1 0 0 0 Schwenk 2000, Whitlock Amphisbaenia 0 0 1 0 0 1 1 0 0 0 and Richman 2013, Zaher and Rieppel 1999. Leptotyphlopidae 0 0 1 0 0 0 1 0 0 0 Typhlopidae 0 0 1 0 0 0 1 0 0 0 Pythonidae 0 0 1 0 0 1 0 0 0 0 Boidae 0 0 1 0 0 1 0 0 0 0 Lamprophiidae 0 0 1 0 0 1 0 0 0 0 Colubridae 0 0 1 0 0 1 0 0 0 0 Elapidae 0 0 1 0 0 1 0 0 0 0 Viperidae 0 0 1 0 0 1 0 0 0 0 AVES ------Louchart and Viriot 2011.

42

Supplemental Table 2.3. Tooth Replacement and Attachment in Mammalia and Dinosauria.

REPLACEMENT ATTACHMENT References Mp. Dp. Pp. Elo. Th. Acr. Pl. Ank. Ped. Con. MAMMALIA Rodentia 0 0 0 1 1 0 0 0 0 0 Chiroptera 0 1 0 0 1 0 0 0 0 0 Soricidae 0 1 0 0 1 0 0 0 0 0 Primates 0 1 0 0 1 0 0 0 0 0 Carnivora 0 1 0 0 1 0 0 0 0 0 Artiodactyla 0 1 0 0 1 0 0 0 0 0 Dirptodontia 1 1 0 0 1 0 0 0 0 0 Lagomorpha 0 0 0 1 1 0 0 0 0 0 Didelphimorphia 1 1 0 0 1 0 0 0 0 0 Cetacea 1 0 0 0 1 0 0 0 0 0 Berkovitz and Shellis 2018, Ungar 2010, Dasyuromorphia 0 1 0 0 1 0 0 0 0 0 Whitlock and Richman Afrosoricida 0 1 0 0 1 0 0 0 0 0 2013. Erinaceomorpha 0 1 0 0 1 0 0 0 0 0 Cingulata 0 1 0 0 1 0 0 0 0 0 Peramelemorphia 0 1 0 0 1 0 0 0 0 0 Scandentia 0 1 0 0 1 0 0 0 0 0 Perissodactyla 0 1 0 0 1 0 0 0 0 0 Macroscelidea 0 1 0 0 1 0 0 0 0 0 Pilosa 0 0 0 1 1 0 0 0 0 0 Monotremata ------Proboscidea 0 0 1 0 1 0 0 0 0 0 DINOSAURIA Ornithischia 0 0 1 0 1 0 0 0 0 0 LeBlanc et al. 2017, Saurischia 0 0 1 0 1 0 0 0 0 0 Sander 1997, Serano 1997.