Phylogeny and Relationships of Taeniodonta, an Enigmatic Order of Eutherian Mammals (Paleogene, North America)

Home , Afrotheria, Cimolesta, Cimolestes, Ectoganus, Kennalestes, Metacheiromys

Thesis

Presented in Partial Fulfillment of the Requirements for the Degree Master of Science in the Graduate School of The Ohio State University

Deborah Lynn Weinstein

Graduate Program in Evolution, Ecology, and Organismal Biology

The Ohio State University

2009

Thesis Committee:

John Hunter, Advisor

William Ausich

John Wenzel

Deborah Lynn Weinstein

2009

Abstract

The Taeniodonta is group of eutherian mammals from the Paleogene of North

America, whose exact place in eutherian phylogeny is uncertain. Taeniodonts evolved rapidly in the Paleocene to achieve, in some forms, large body size, hypselodont (i.e., evergrowing) canine and postcanine teeth, and peculiar patterns of tooth wear. Eleven genera of taeniodonts occur in two main subclades, recognized at the level of families or subfamilies depending on author, the Conoryctidae and the Stylinodontidae. The conoryctids were smaller, probably insectivorous or omnivorous, and retained a larger number of primitive characters than did the stylinodontids. The stylinodontids were larger than the conoryctids, possessed massive canines, and exhibit a trend toward hypselodonty of the canines and molars. Prior to this study, there has not been a comprehensive phylogeny of all of the currently recognized genera of taeniodonts. In

Chapter 1, I review the history of research on the systematics, evolution, and paleobiology of the taeniodonts, with emphasis on recent discoveries of transitional forms. I identify the major unresolved phylogenetic issues that concern taeniodonts that

I explore in subsequent chapters including the internal relations among taeniodonts,

monophyletic versus diphyletic origins, ancestry, and the status of the taeniodonts as either stem or crown eutherians.

In Chapter 2, Internal Relations Among the Taeniodonts, I take up the relations of genera of taeniodonts to one another and overall biogeographic history. I performed cladistic analyses using NONA and Winclada to determine the evolutionary relations of

the known genera of the Taeniodonta. Two clades are well supported, a clade consisting of the conoryctids exclusive of Onychodectes (i.e., Conoryctella, Conoryctes, and

Huerfanodon) and a clade consisting of all the known stylinodontids (Wortmania,

Schochia, Psittacotherium, Ectoganus, and Stylinodon). Stratocladistic analysis, which takes into account the temporal sequence of taxa in the fossil record, supports the results of phylogenetic analysis using morphological characters alone. Phylogenetic, stratigraphic, and geographic data contribute to reconstructing the biogeographical history of the taeniodonts. In these analyses, the Late Cretaceous taeniodont

Schowalteria occupies the most basal position in taeniodont phylogeny, pre‐dating an inferred conoryctid–stylinodontid split in the early Paleocene.

In Chapter 3, External Relations of the Taeniodonts, I investigate the higher‐level relations and ancestry of the taeniodonts as well as the monophyletic or diphyletic origins. I describe a newly discovered fossil lower jaw from the early Paleocene of North

Dakota that I interpret to be the lower jaw of Alveugena, a taxon previously argued to be a transitional form between cimolestids and the taeniodonts. This discovery contributes new characters and facilitates study of the relations between taeniodonts

iii

and basal cimolestans. This study corroborates the hypothesis that Alveugena is the sister taxon of the taeniodonts. Contrary to earlier analyses, the Late Cretaceous–

Paleocene Cimolestes is found to be the sister taxon of Alveugena+Taeniodonta rather than the Paleocene Procerberus, thought by some to be closer to the taeniodonts.

Stratocladistics provided additional support for the conclusions of the analyses of morphology alone and implicate Cimolestes as a possible ancestor to Alveugena and the taeniodonts. Further cladistic analyses including taeniodonts, Late Cretaceous stem eutherians, and Paleogene representatives of crown eutherian clades determine the higher‐level relations of taeniodonts. This study shows that the taeniodonts lack the synapomorphies that link crown eutherians together.

In the final chapter, Conclusions, I will discuss the overall aspects of the taeniodonts. I will also discuss the evolutionary rates of the canine and molar evolution in the group, as well as rediagnosing the clades.

In conclusion, the taeniodonts are a monophyletic group of stem eutherian mammals whose closest known sister group is Alveugena, and most likely Cimolestes gave rise to the Alveugena and the taeniodonts.

iv Acknowledgements

I first thank Dr. John Hunter, for getting me started with this project and being there through all of it, and for helpful discussions on thesis matters. I would also like to thank my intrepid committee members, John Wenzel for his help with the cladistics found within and Bill Ausich for help with the past and with my future. I would also like to thank Judy Galkin for her tireless efforts as I searched the American Museum of

Natural History collections and Michael Brett‐Surman for his help at the National

Museum of Natural History. I thank David Fox and especially Jonathan Marcot for help using their program StrataPhy. Finally, I would like to thank the Ohio State University

Department of Evolution, Ecology, and Organismal Biology and a University Fellowship and Teaching Associateship that have supported my work.

v Vita

June 2003………………..………………………………….Revere High School

2007…………………………………………………………….B.A. Biology & Evolutionary Biology, Case

Western Reserve University

2007–2008…………………………………………………..University Fellowship, Ohio State University

2008–2009…………………………………………………..Graduate Teaching Associate,

Department of Evolution, Ecology and

Organismal Biology, Ohio State University

Publications

Croft, D.A and D. Weinstein. 2008. The first application of the mesowear method to endemic South American ungulates (Notoungulata). Palaeogeography, Palaeoclimatology, Palaeoecology 269: 103–114.

Fields of Study

Major Field: Evolution, Ecology, and Organismal Biology

vi Table of Contents

Abstract………………………………………………………………………………………………………………………….ii

Acknowledgements………………………………………………………………………………………………………..v

Vita………………………………………………………………………………………………………………………………..vi

Table of Contents………………………………………………………………………………………………………….vii

List of Tables………………………………………………………………………………………………………………..viii

List of Figures…………………………………………………………………………………………………………………ix

Chapter 1: Taeniodont Overview and the History of Their Classification…………………………1

Chapter 2: Internal Structure of Taeniodont Phylogeny……………..…………………….………….20

Chapter 3: Taeniodonta's Placement in the Phylogeny of Eutherian Mammals ……………32

Chapter 4: Conclusions…………………………………………………………………………………………………45

References……………………………………………………………………………………………………………………54

Appendix A: Description of the lower jaw of Alveugena……………………………………………….59

Appendix B: Characters and Matrices…………………………………………………………………………..66

vii List of Tables

Table 1. Literature used to code characters for the genera and species in the study..….21

Table 2. List of biogeographic placement of taeniodont specimens..……………………………27

viii List of Figures

Figure 1. Schoch's (1986) reconstruction of Onychodectes……………….…………………………….1

Figure 2. Schoch's (1986) reconstruction of Stylinodon…………………………………………………..1

Figure 3. Skulls of Conoryctes (1) and Psittacotherium (2) from Matthew (1937)……………2

Figure 4. Stylinodon right manus from Matthew (1937)….……………………………………………..3

Figure 5. Known chronostratigraphic distribution of the taeniodonts.………...…………………5

Figure 6. Plot of known Taeniodonta localities…..…………………………………………………………..6

Figure 7. Schoch’s (1986) phylogeny of taeniodont genera with families also shown.....10

Figure 8. Lucas and Williamson’s (1993) phylogeny………………………………………………………12

Figure 9. Eberle’s (1999) most parsimonious phylogeny……………………………………………….15

Figure 10. Internal structure tree from cladistic analysis.....………………………………………….23

Figure 11. Internal structure trees from stratocladistic analysis…..……………………………….25

Figure 12. General movements of the taeniodonts based on biogeographical data..……28

Figure 13. Biographically optimized internal tree from cladistic analysis.……………………..30

Figure 14. Cimolestid relations of tree from cladistic analysis………………….…………………..33

Figure 15. External relations tree from cladistic analysis……………………..………..………..……35

Figure 16. External relations tree from stratocladistic analysis…….….……………………………40

Figure 17. External relations among stem Eutheria from cladistic analysis…………………..42

ix Figure 18. External relations among crown Eutheria from cladistic analysis…………………43

Figure 19. Known taeniodont stratigraphic distributions with phylogeny.…………………….46

Figure 20. Patterns of character divergence in canines and first molars.………………………49

Figure 21. Alveugena lower jaw in buccal, lingual, and occlusal view……………………………61

x Chapter 1: Taeniodont Overview and the History of Their Classification

Interest in the origin of placental mammals has led to the discovery of many mammalian taxa. Although several of these groups do not lead to extant mammals, they are part of the overall biotic environment in which extant placental mammals

originated. One of the most interesting of these groups is the Taeniodonta, which has been viewed variously in the Linnean hierarchy as either an order (Rose, 2006;

Schoch, 1986) or a suborder (McKenna and Bell, 1997) of eutherian mammals. In this study, Taeniodonta will be viewed as an order, which is the more common practice.

Taeniodonta ranged from small, insectivore‐like mammals (Figure 1) to large, pig‐like animals (Figure 2). The group includes several genera: Onychodectes Cope,

Figure 2. Schoch's (1986) Figure 1. Schoch's (1986) reconstruction of reconstruction of Stylinodon. Onychodectes.

1 1888a, Wortmania Hay, 1899, Conoryctes Cope, 1881, Conoryctella Gazin, 1939,

Huerfanodon Schoch and Lucas 1981a, Psittacotherium Cope 1882a, Ectoganus Cope,

1874, Stylinodon Marsh, 1874, Schochia Lucas and Williamson, 1993, and Schowalteria

R. C. Fox and Naylor, 2003. Robert Schoch, who published a comprehensive monograph on Taeniodonta in 1986, classified the known genera of taeniodonts into two families, the Stylinodontidae and the Conoryctidae. The conoryctid genera that were known at the time of Schoch (1986) were Onychodectes, Conoryctes, Conoryctella, and

1 2

Figure 3. Skulls of Conoryctes (1) and Psittacotherium (2) from Matthew (1937). Evident here are the large, oddly shaped canines and more robust skull of the stylinodontids. Images not to scale.

Huerfanodon. Conoryctidae includes small animals that retained more primitive characteristics than the Stylinodontidae. All taeniodonts, however, have some degree of

2 tooth crown hypsodonty. Conoryctes (Figure 3.1) as well as Onychodectes (Figure 1) are representative of this group. The Stylinodontidae at the time of Schoch (1986) included

Wortmania, Psittacotherium, Ectoganus, and Stylinodon. Other than the tooth crown hypsodonty that the stylinodontids share with the conoryctids, this group is has more

derived characters including increased body size, larger canines, and much larger and more robust skulls, for example Psittacotherium (Figure 3.2) Stylinodon (Figure 2).

Stylinodontids are also characterized by enlarged, recurved, and laterally compressed

Figure 4. Stylinodon right manus from Matthew (1937).

claws on their hands (Figure 4). Schowalteria and Schochia were both discovered since

Schoch's monograph. R. C. Fox and Naylor (2003) initially placed Schowalteria in the

3 Stylinodontidae. Lucas and Williamson (1993) placed Schochia as the sister group to the

stylinodontids.

Taeniodonta first appeared during the Late Cretaceous (R. C. Fox and Naylor,

2003) and became extinct in the middle Eocene (Turnbull, 2004). Schowalteria of the

Late Cretaceous of Alberta, Canada was the earliest taeniodont (R. C. Fox and Naylor,

2003). Stylinodon was the only genus that persisted into the Uintan North American

Land Mammal Age (NALMA) (middle Eocene) (Schoch, 1986). Psittacotherium was the first truly large‐bodied taeniodont. Schoch (1986) estimated the body mass

Psittacotherium at 35.6–71.2 kg, based on skull and body length, of, which was larger than Wortmania (the largest Puercan taeniodont) at 14.4‐28.8 kg. The Taeniodonta has been considered part of the great diversification of mammals following the mass extinction at the end of the Cretaceous, which ended the reign of the dinosaurs (R. C.

Fox and Naylor, 2003). The diversification of the Taeniodonta is most likely still linked to

this great mammal diversification because most of the genera (all but Schowalteria) do not appear until the Paleocene. The chronological distribution is shown in Figure 5.

4 Figure 5. Known chronostratigraphic distribution of the taeniodonts. Boxes show the differing classification of Schoch (1986) and McKenna and Bell (1997).

Taeniodonta are also restricted in their spatial distribution. So far, taeniodonts

have only been found in North America, centered in the western United States and

Canada (Figure 6). The discovery of Schowalteria clemensi R. C. Fox and Naylor, 2003

changed many ideas about the spatial extent of Taeniodonta, which before had been

known as far north as North Dakota. Researchers found this species in Alberta, Canada,

which is several hundred kilometers north of the northern limit of previous taeniodonts.

It is also possible that the origin of this group occurred much farther north than thought

(R. C. Fox and Naylor, 2003). 5 Figure 6. Plot of known Taeniodonta localities. Created using data downloaded from the Paleobiology Database on April 1, 2008, using the group name 'Taeniodonta' and the following parameters: Country = United States; Country = Canada.

Mammalian paleontologists study Taeniodonta for many reasons. First, these species have wear on their teeth that differs from that of any other group of mammals, with pronounced cusp wear over the entire occlusal surface leading to complete destruction of the chewing surface. This wear pattern does not only make them difficult

to relate to other animals, but it can commonly lead to lack of information due to removal of all dental surfaces. Second, this order was one of the first mammal groups to reach large body size (Psittacotherium at 35.6‐71.2 kg; Schoch, 1986). These changes in tooth morphology and size happened over a short amount of evolutionary time, which led Patterson (1949) to infer a rapid rate of evolutionary change in this group.

6 The evolutionary rate that Patterson (1949) noted is in the Stylinodontidae, because there were major changes in very little time. The taeniodonts are found over a

33‐million‐year range, and there are nine genera. However, during this time the

Stylinodontidae changed greatly in form. There were huge changes in the feet, with

trends toward digging appendages with large claws. Also, the canines of the group reached a morphology where they have both a cutting edge and a grinding surface in the back. This morphology was achieved between Wortmania and Psittacotherium, which occurred within the early Paleocene, in about 5‐6 million years. There was a swift

change in molar evolution toward the terminal end of the group that also lasted 5‐6 million years. Although the actual rates of evolution have not been calculated, the form of the stylinodontids changed drastically in just a few million years (Patterson, 1949).

Biogeography

Taeniodonta has only been found in North America. Though the western interior is where the majority of the specimens are found, there seems to have been an expansion of geographic range in later species. Ectoganus made it to the east coast

(Aquia Formation, Maryland and Williamsburg Formation, South Carolina) and

Psittacotherium and Stylinodon dispersed to the southern coast of Texas (Black Peaks

Formation and Devil's Graveyard Formation, respectively; see Figure 6). The farthest north that a taeniodont has ever been found is Alberta, Canada, and that was

Schowalteria, the earliest known taeniodont. Because this early taeniodont appears in

7 Alberta, Canada, it seems plausible that Taeniodonta originated further north and then migrated south to where most taeniodonts have been found. This hypothesis is supported by fossil evidence from other groups, such as neoplagiaulacid multituberculates and condylarths, which lived concurrently and arguably originated in

Canada and migrated south (R. C. Fox and Naylor, 2003) The earliest Cenozoic

specimens of Taeniodonta appear in the northern latitudes. Onychodectes occurred in

Montana and then relatively soon after found in New Mexico. These lines of evidence lead me to infer Taeniodonta originated in the north, moved down to separate northern and southern basin groups and then diversified from there.

History of Taeniodont Internal Relations

The first classification of this group was completed in the late nineteenth century, when Wortman (1896, 1897) divided this group into two families: the

Conoryctidae and the Stylinodontidae. Originally these two families were placed in the suborder Ganodonta, which contained only taeniodonts that had been discovered by

this time. This group name would later be changed due to Cope's description of the rodent‐like Taeniodonta. Conoryctidae consisted only of Onychodectes and Conoryctes, whereas Stylinodontidae consisted of Hemiganus Cope, 1882a (now considered to be a junior synonym of Wortmania), Psittacotherium, Calamodon Cope, 1874 (now considered to be a junior synonym of Ectoganus), and Stylinodon (see Wortman, 1896,

1897).

8 Matthew (1937) described the group as an evolutionary series from

Onychodectes to Stylinodon, except that Conoryctes was an off‐shoot that had become divergent from the rest of the group. This series was based on adaptations of anatomy that had changed through a changing environment. He described the group as one

family (Stylinodontidae) with four subfamilies: Onychodectinae (Onychodectes),

Conoryctinae (Conoryctes), Psittacotheriinae (Wortmania, Psittacotherium, Ectoganus

and other synonyms), and Stylinodontinae (Stylinodon).

Schoch's (1986) monograph used Wortman's (1896, 1897) families, adding taxa

discovered more recently as well as correcting naming problems that had arisen in the

ninety years between Wortman’s and Schoch’s studies. Schoch’s phylogeny based on his

descriptions of the taeniodonts known at the time is shown in Figure 7. His phylogeny followed the naming of the two families. It uses an unknown “Taeniodont Ancestor” as the root of the analysis (Schoch, 1986). Schoch (1986) performed the analysis by hand, without the aid of computer programming, and did not use an explicit outgroup to polarize characters in the analysis. Furthermore, this "Taeniodont Ancestor" is a hypothetical construct with the most basal characters that could have been present in an ancestor to the group.

9 Figure 7. Schoch’s (1986) phylogeny of taeniodont genera with families also shown.

McKenna and Bell (1997) changed the familial naming, agreeing with Matthew

(1937), but reduced his number of subfamilies to two: Conoryctinae and

Stylinodontinae, which resemble Schoch’s original families. The only difference between

McKenna and Bell's subfamilies and Schoch's (1986) families is that there is no subfamily distinction for Onychodectes and Conoryctella (McKenna and Bell, 1997).

A new discovery since the last monograph of Taeniodonta was Schochia sullivani

Lucas and Williamson, 1993, which the authors state has characters of both taeniodont families and their suspected ancestors. This small taeniodont is from the middle Puercan

Pu2 interval zone, in the same rock layer as the genera Onychodectes and Wortmania. It has premolars that are more derived than premolars in the Conoryctidae, but molar characters that are less derived. The researchers described the premolars as resembling

10 those of Stylinodontidae in structure, whereas they described the molars as resembling those of the primitive group that Schoch claimed to be ancestral to the taeniodonts

(Lucas and Williamson, 1993).

Lucas and Williamson (1993) argued that Schoch (1986) should not have made the existence of crown hypsodonty a defining characteristic for either family because it was a synapomorphy for the entire order. Of course, a higher degree of hypsodonty could be used as characteristic of Stylinodontinae, because all species express it, which is how it is approached in the following studies. Also, Lucas and Williamson protested the use of Schoch’s premolar characteristics, which are shared with a possible ancestral

group Cimolestidae (including Cimolestes Marsh, 1889, and Procerberus Sloan and Van

Valen, 1965) (Lucas and Williamson, 1993). This problem is resolved in this study by using characters from both Schoch's (1986) analysis and an analysis of that ancestral group (the cimolestids).

Lucas and Williamson (1993) added Schochia to the list of taeniodonts and generated a phylogeny, based on a few morphological characteristics, which placed S. sullivani between Schoch’s two families (Figure 8). This phylogeny was only based on five morphological characters and only used four taeniodont genera and a combination of the rest as "Stylinodontidae". The five morphological characters were placed by hand on the phylogeny without the help of computer programming. Also, the lack of the individual genera within the stylinodontids may have steered the analysis toward placement of Schochia as the sister taxa to the group.

11 Figure 8. Lucas and Williamson’s (1993) phylogeny.

Lucas and Williamson (1993) explained that this classification of Schochia

between the two families would complicate the purported molar evolution in taeniodonts. Because the molars of S. sullivani are more primitive than other taeniodonts, the teeth must have returned to a more primitive state (termed a

“reversal” in character state) for this phylogeny to be correct. They asserted that this reversal is possible although unlikely. Because Psittacotherium also has these more primitive teeth, there would have actually been two reversals in the molar evolution of the taeniodonts, according to these scientists’ phylogeny (Figure 8). Lucas and

Williamson (1993) also proposed that S. sullivani could be part of a sister taxon that has convergent premolar characteristics.

12 Although a phylogeny has not been constructed that includes the newly discovered Late Cretaceous taeniodont Schowalteria, its discoverers discussed its many shared characteristics with the later Stylinodontidae. Schowalteria is known from a fragmentary skull preserving part of the dentition. However, its premolar and molar characteristics are much more similar to earlier eutherian mammals when compared to

a contemporary cimolestid (R. C. Fox and Naylor, 2003).

History of Taeniodont Higher Level Classification and Eutherian Relationships

Over the years, paleontologists have allied the taeniodonts with many different groups of early mammals based on appendage and tooth morphology. Cope (1888b) placed the taeniodonts, along with the tillodonts, as part of the ancestral group of

Rodentia because of his description of their hypertrophied canines of Psittacotherium and Ectoganus as rodent‐like incisors. Marsh (1875) originally placed the stylinodontids with the tillodonts, due to similar tooth (though "small canines") and appendage morphology. Wortman (1896, 1897) placed them as the origin of the Edentata (now known as Xenartha, e.g. armadillos, sloths) in North America, whereas Matthew (1937) considered them as possibly outlying edentates. Aligning Taeniodonta with the Edentata is understandable in light of the shared adaptations for digging of the Stylinodontidae, but the resemblance is probably due to evolutionary convergence. Simpson (1931a)

dispelled the thought of taeniodont relationships with Edentates by discussing each

13 factor and determining that convergence was probably playing a strong role and that the taeniodonts probably had their origin in one of the early insectivore groups.

During the late 1960s, Simpson's (1931) idea was finally examined in detail, and an ancestral group was chosen. McKenna (1969) and Lillegraven (1969) discussed the

possibility of the taeniodonts having arisen from Procerberus, an insectivore closely related to, and possible descended from, Cimolestes. Through the past forty years, the classification of these two animals has changed. Now, they are considered solidly cimolestids, but their placement as the ancestral group to the taeniodonts is still under

debate.

A discovery relevant to this question was Alveugena carbonensis Eberle, 1999, claimed to be the largest species of the family Cimolestidae by the author. This species follows a morphological gradient, especially in size and dental characteristics, that seems to flow from Cimolestes through Procerberus and finally into Alveugena in the

cimolestid lineage. Alveugena was placed with the cimolestids because it lacks hypsodonty on its cheek teeth, otherwise it would have been placed with the taeniodonts. Temporally, it falls directly between Procerberus grandis Middleton and

Dewar, 2004 and Onychodectes, which were latest early Puercan (Pu1) and middle

Puercan (Pu2), respectively. Alveugena also has features that would place it between these two on the morphological gradient between cimolestids and the early taeniodonts. For example, there is an increase in size and robustness of the lower fourth premolar and molars and a transverse narrowing in the molars. Eberle (1999), seeing

14 these trends from Cimolestes all the way to Onychodectes, inferred that there was a diet shift from the more insectivorous Cimolestes and Procerberus formicarum Sloan and Van

Valen, 1965 to a more omnivorous Alveugena and Onychodectes. There was also an increase in body size, evidence by an increase in tooth size, throughout this lineage.

Using these characters and others, Eberle (1999) performed a cladistic analysis, resulting

Figure 9. Eberle’s (1999) most parsimonious phylogeny.

in the tree in Figure 9. This agrees with Eberle's claim that Alveugena falls directly between the two Procerberus species and Onychodectes. She also argued that there are no autapomorphies precluding Alveugena as the ancestor of Onychodectes, and the

15 timing is close to that predicted by the hypothesis of an evolutionary transition from

Procerberus, through Alveugena, to Onychodectes. There was only upper dentition available from this species, so relationships were determined only from upper tooth characteristics (Eberle, 1999). However, a newly discovered specimen from the

Paleocene of North Dakota will increase the number of known characters: a lower jaw most probably of Alveugena, which is described in Appendix A.

Eberle acknowledged that the phylogeny had several problems. First, a reversal in characters is required if Alveugena is the descendant of one of the Procerberus species. The upper fourth premolars in Procerberus possess a metacone, whereas the metacone of this tooth in Onychodectes or Alveugena is either reduced or nonexistent, respectively. This either suggests a character reversal, or that the Procerberus species are actually a sister group to the Alveugena and neither is an ancestor of the other

(Eberle, 1999). The analysis was also done using a Wagner tree (which is one of the simpler cladistic methods, but not the most robust analysis), and it was done with only fifteen characters. Finally, only one taeniodont was chosen in this analysis, whereas more information could be learned from having all the genera included (though then more characters would have been needed). The one taeniodont chosen was the most earliest that was known at the time, but an earlier animal, Schowalteria, has since been found.

Another place of debate for the taeniodonts is the monophyly of the group.

Because the two subgroups are so different, many scientists have found it hard to

16 believe that they have one origin (see above distinctions of stylinodontids as rodents or tillodonts while the conoryctids were allied with early insectivores). Schowalteria, with its older age and supposed stylinodontid characters seems to make it impossible for the taeniodonts to be monophyletic (R. C. Fox and Naylor, 2003). However, no cladistic

analysis including Schowalteria has been completed, until the studies in Chapters 2 and

The relationships of taeniodonts to the rest of Mammalia will be examined in

Chapter 3. Not only will I test the placement of Alveugena as a sister group to the

Taeniodonts, but I will also consider the placement of the taeniodonts among other

lineages of early mammals and the modern clades (Afrotheria, Euarchontaglires, etc).

These and the monophyly of the group will be examined and discussed in that chapter.

Stratocladistics

Stratocladistics is a relatively new method that uses stratigraphic information along with morphological data to interpret the phylogenies of ancient groups.

Reconstructing phylogenies stratigraphically can show a few things that simple cladistics can miss, such as the sequence in which the characters appeared in time (Benton, 1995).

Stratocladistics uses character debt to interpret tree length, as in parsimonious cladistic analyses, as well as stratigraphic debt for placing a taxon in an improper order compared to the fossil record. Stratigraphic debt increases with the length of hypothetical ghost lineages that connect taxa. It creates actual phylogeny‐specific

17 hypotheses of evolution, and not just parsimonious trees (D. L. Fox et al., 1999). Because of this, stratocladistics could be a great asset to the paleontological community and to the evolutionary biology community. Recently, a program that can do such analyses has become available, StratPhy, which will make this new tool in the paleontologists' kit

easier to use and understand (Marcot and D. L. Fox, 2008). This new type of analysis will help determine whether issues with stratigraphic placement, such as Schowalteria's early age, would really be problems within a phylogenetic analysis. Also, this type of analysis is another way of arriving at conclusions, and adds to the amount of information available in each analysis.

Summary of Unresolved Questions Regarding the Taeniodonta

The evolution of the taeniodonts has led to debates about the way that the group should be structured. Between Schoch's (1986) and McKenna and Bell's (1997) categorizations, there is dispute about the placement of Onychodectes and Conoryctella, either they both belong with the conoryctids or neither of them do. Similarly, the placement of Schochia as the sister group to the stylinodontids has complicated matters of taeniodont evolution (Lucas and Williamson, 1993). Finally, Schowalteria has been placed with the stylinodontids, although conjectured and not analyzed through cladistics. Here, in Chapter 2, the first computer aided parsimony analysis of this group has been completed, with all known taeniodonts. This analysis takes into consideration characters from both the taeniodonts as well as their purported ancestral group, the

18 cimolestids. I will test the placement of the recently discovered taxa (Schochia and

Schowalteria) and also test whether the conoryctid and stylinodontid groupings withstand scrutiny.

The placement of the taeniodonts within the evolution of mammals has also been studied and debated for over a hundred years. The current thinking is that the

taeniodonts belong as sisters to the cimolestids (including Cimolestes, Procerberus, and

Alveugena) and that Alveugena is the closest sister taxa followed by Procerberus. The monophyly of the group also has been debated, because Schowalteria's early age and placement as a stylinodontid separates the group in two, due to the ghost lineages that

would be required for the conoryctids to have evolved from an ancestor earlier than

Schowalteria. Finally, the placement of the taeniodonts as stem or crown eutherians has never really been studied. In Chapter 3, a computer aided parsimony analysis has been done, using outgroups from the cimolestids and further outgroups to determine whether the cimolestids are the true sister group and whether the taeniodonts are a monophyletic clade. Finally, an analysis placing the taeniodonts into their context on the eutherian mammal tree is also presented.

19 Chapter 2: Internal Structure of Taeniodont Phylogeny

The debate about the phylogenetic arrangement of the taeniodonts into families or subfamilies has persisted since their early discovery. New taxa seemed to add to the confusion instead of answering questions, but a rigorous cladistic analysis of the entire known group has not been accomplished previously. The last phylogenetic analysis

(Schoch, 1986) at the generic level was done by hand, not computer‐aided, and was completed more than twenty years ago. Since then several new species and new specimens of previously known species have been found and described. In the analyses that follow, the entirety of known taeniodonts is finally considered as a whole.

Cladistic Perspective

The first analysis run was a parsimony analysis. The characters used were augmented and modified from two main sources: Eberle's (1999) analysis of the placement of Alveugena and Schoch's (1986) monographic treatment of the taeniodonts. The characters from Schoch's (1986) monograph had not been coded, and

the Eberle (1999) characters were expanded to apply to all of the taeniodonts (Eberle,

1999, coded only Onychodectes among the taeniodonts as a representative). Any overlapping characters were removed. This procedure left thirty‐seven characters to be coded for eleven taxa, including all known Taeniodonta. Characters were coded using

20 Table 1. Literature used to code characters for the genera and species in the study.

the literature (see Table 1), the description of the lower dental morphology of

Alveugena (Appendix A), and observations of specimens at the American Museum of

Natural History (AMNH), New York, and the National Museum of Natural History

(NMNH), Washington.

The goal of cladistic analysis in general is to find the shortest length tree

available from a set of taxa and characters. There are many ways employed by many different programs currently in use by systematic scientists. NONA, the program used in all analyses here, first creates many Wagner tree replicates using random addition of taxa, and then swaps branches using tree‐bisection reconnection (TBR) (Goloboff, 1993).

A Wagner tree is built by starting with one taxon and then adding taxa in such a way that the fewest additional evolutionary steps will be added (Farris, 1970). Doing

21 different replicates of the Wagner tree will build trees of different topologies based on

the order in which the taxa are added. TBR is a branch swapping mechanism that cuts the Wagner tree into two subtrees and then reattaches one segment to the other at any point on the other subtree. In this way, any two branches on the tree could be connected (Siddall, 2002). NONA is also capable of finding the Bremer support values for any particular most parsimonious tree that it has found. Bremer supports show how many steps longer a tree has to be before the clade in question is lost (Wenzel, 2002).

One of the optional interfaces for NONA, Winclada, was also used in this analysis, and has the ability to calculate a consistency index (CI) and a retention index (RI), homoplasy indicators, for the trees produced by NONA. The CI is based on the number of additional

evolutionary steps taken in a character. This statistic shows the amount of homoplasy in a tree. The RI, based on worse case scenarios, determines how far the character evolution is away from the most number of steps possible. This statistic also indicates the amount of homoplasy in the tree (Wenzel, 2002).

22 Figure 10. Internal structure tree from cladistic analysis. Single most parsimonious tree found in NONA, with a length of 123, CI of .55, and RI of .68. Bremer supports from NONA added, and outgroups Cimolestes, P. formicarum, P. grandis, and Alveugena removed for space.

Results

The results of the analysis are summarized in Figure 10. The results also show that one clade containing Conoryctella, Conoryctes, and Huerfanodon and another containing Wortmania, Schochia, Psittacotherium, Ectoganus, and Stylinodon are both well supported with a Bremer support of 3 or more. It is also interesting that the clade

that splits the taeniodonts into the two subclades is also well‐supported (Wenzel, 2002).

There are several characters holding these groups together. For the taeniodonts, although they are not as well supported as the conoryctid and stylinodontid clades, they

23 are still bound by several molar characters including lack of cingula/ids on upper and lower molars, trigonids subequal in size to talonids, and the molar hypsodonty for which they are so well known (Schoch, 1986). The conoryctids, which have been suggested as a paraphyletic group (Lucas and Williamson, 1993), are here well supported with several

characters holding them together. The deep root of the canine and p4 being submolariform are nonmolar characters holding them together whereas the upper molar characters include a large metacone, absent conules, and a mesostyle on M1 and

M2 that is not absent as it is in the other taeniodonts (and increased in size through the evolution of the group). Not only are the conoryctids well‐supported as a group, but the cladistic analysis can point us to characters that hold the clade separate from the remainder of the taeniodonts. There are several characters holding the stylinodontids

together, including single upper and lower incisors, oblique lower premolars, a well‐ developed p4 talonid heel, incipient P4 stylocone, bilophodont lower molars and transverse upper molars. The support of this group is based on the characters that hold it together, and these characters can point us in the direction of the evolution of the

group.

This analysis shows that there is a monophyletic group of conoryctids, although lacking Onychodectes and a monophyletic group of stylinodontids. Schowalteria falls out as the most basal taeniodont, and not as a cause for paraphyly in the group.

24 Stratigraphical Perspective

In paleontology, fossils occur in a stratigraphic sequence. Although this stratigraphy is not commonly used in phylogenetic analyses, it can help to determine the plausibility of not only a cladistic analysis in light of stratigraphic sequence, but also helps to determine ancestor‐descendent relationships. For my analysis, I used StrataPhy,

a new program that uses Wagner trees and TBR as its cladistic analysis. Once trees are found, it can place taxa as ancestors based on stratigraphic sequence and lack of autapomorphies in the ancestral taxa (Marcot and D. L. Fox, 2008).

The resulting most parsimonious trees look strikingly similar to the tree produced by cladistic analysis (Figure 11). These two trees were found using both the exhaustive and heuristic searches for ancestors. The only differences stem from the

Figure 11. Internal structure trees from stratocladistic analysis. Most parsimonious trees found in StrataPhy. Horizontal taxa represent ancestral placement.

25 ability of StrataPhy to place ancestors. In this analysis, in both trees, Cimolestes is placed as the ancestor to a clade of Alveugena+Taeniodonta and a clade of the Procerberus species. Also in both trees, Schochia is placed as the ancestor to the remainder of the stylinodontids (Psittacotherium, Ectoganus, and Stylinodon). The only difference

between the two most parsimonious trees is that one places Procerberus formicarum as

the ancestor to Procerberus grandis, whereas the other leaves them only as sister taxa.

This distinction is outside of the classification and study of the taeniodonts; and therefore, is not a problem for the internal structure of the taeniodonts.

This is another line of evidence pointing to the plausibility of the phylogeny that has been found in this analysis. Agreement between stratigraphic information and cladistic analysis shows even more support for the clades within the analysis as well as the tree as a whole.

Biogeographical Perspective

Biogeography is another tool that we have, assuming that the fossils of terrestrial animals are found in the same local area of at least the same region as the original animals (Behrenmeyers et al., 1992: 80). Although there are pieces of the fossil record missing, we have the occurrences of the fossils and can infer their time of existence. Therefore, we can conjecture how the evolution and diversification of a group may have unfolded to account for the distribution of taxa. Table 2 shows the approximate dates and locations of known occurrences of the taeniodonts.

26 Table 2. List of biogeographic placement of taeniodont specimens. Found using the Paleobiology Database, dates are ranges shown reflecting individual authors' approximation of date. Multiple specimens from the same approximated time are not shown to conserve space.

27 Figure 12. General movements of the taeniodonts based on biogeographical data. Dots are taxa, straight arrows are movement of taxa with which it shares a color and blue curved arrows are transitions.

The geographic distribution of fossil occurrences when connected to a well‐

supported phylogeny can be informative about biogeographic history. Stratigraphic

information is also important in paleobiogeography because dates are vital to

understanding the movement of organisms on the landscape. The story that this

information tells is a fascinating one of dispersing mammals and places of origin for

various clades. Schowalteria moved southward from its Canadian location until reaching

Montana, where Onychodectes is found earliest. A group of Onychodectes‐grade

28 animals moved south toward New Mexico while another group stayed behind in the north. The northern group of Onychodectes may have evolved into Conoryctella, which then moved south to New Mexico to evolve into Conoryctes and then Huerfanodon. All the conoryctids eventually moved back north (perhaps changing habitats led the

conoryctids to be better suited to northern latitudes). Meanwhile, the Onychodectes group that had moved south was branching into the stylinodontids, starting with

Wortmania and quickly Schochia, which are found in New Mexico. Schochia then most likely moved north to where Psittacotherium was first found, and eventually evolved into Ectoganus and Stylinodon. Psittacotherium moved south again through New Mexico all the way into Texas. Ectoganus and Stylinodon groups seem to have split, both leaving one group in the northern states and an Ectoganus group made it all the way to the East

Coast while a Stylinodon group made it to southern Texas.

The only part of this story that disagrees with the above analyses is the transition of Schowalteria to Onychodectes transition. In the cladistic analyses, Onychodectes diverged into just the stylinodontids and has no part in the origin of the conoryctids. The biogeographic evidence seems to point to Onychodectes spawning both families,

because it is in the right place at both of those right times. Evidence for the northern

Onychodectes is not strong, since it was one tooth that may have been misidentified, but for now, it is possible that an Onychodectes‐grade animal was in the northern

latitudes in the correct time frame. If you were to move Onychodectes to a phylogenetic placement to account for this (Figure 13), then you would change the length of the tree

29 Figure 13. Biographically optimized internal tree from cladistic analysis. Modified in Winclada with length 126, CI .53, and RI .67. Outgroups removed as before.

by only three. Without having some other transitional taeniodont, it is hard to determine which of the biogeographical stories is more probable. However, with the phylogeny found in the cladistic analysis, Onychodectes may have just been moving through, and a Schowalteria‐grade taeniodont spawned the conoryctids.

Familial relations of Taeniodonta

Although the levels of distinction will continue to be debated (whether the two major subclades are families versus subfamilies), it seems clear that there are two main groups of Taeniodonta. In these analyses, the conoryctids are solidly Conoryctella,

Conoryctes, and Huerfanodon. This result opposes McKenna and Bell's (1997) removal of

30 Conoryctella from this group. With these analyses, Conoryctella falls squarely with the other conoryctids. Onychodectes, which was also removed by McKenna and Bell (1997), falls outside of the conoryctids in these analyses. As for the stylinodontids, this group is solidly Wortmania, Schochia, Psittacotherium, Ectoganus, and Stylinodon. The only real counterargument here would be Lucas and Williamson's (1993) placement of Schochia as the sister group to the stylinodontids and not within the group. However, their explanation was not backed with a true cladistic analysis (for instance, they only looked at five characters and grouped all the stylinodontids together), so this new placement is

better supported.

Chapter 3: Taeniodonta's Placement in the Phylogeny of Eutherian Mammals

This chapter focuses on the placement of Taeniodonta within the evolutionary history of mammals. There has been much controversy with the placement of this group, but some definitive answers are beginning to come to light. A new lower jaw of

Alveugena, a possible sister group to the taeniodonts, newly described here, allows in

depth study of the origins of Alveugena within the cimolestids and its connection to the

taeniodonts. I hypothesize that Alveugena will not lose its place as the sister group to

the taeniodonts even with Schowalteria, the more basal taeniodont, and the new information from the lower jaw. The likelihood of Alveugena's origin from a Procerberus line is tested here not only by doing an analysis of known taeniodonts and their purported closest sister taxon, the cimolestids, but also by using other palaeoryctoids. In addition, stratocladistics can determine whether Cimolestes itself could be a possible ancestor to the groups, as has been suggested. While working with the origins of taeniodonts, monophyly can also be tested, by determining whether any outgroup can be added that would break up the clade. Once monophyly has been tested, then the placement of taeniodonts within the diversification of mammals can be established by

studying them in light of stem eutherians, which fall outside of Placentalia and with known crown group eutherians, which lie within Placentalia (Wible et al., 2007).

Cladistic Analyses

The analysis completed in the previous chapter using NONA (Goloboff, 1993).

The thirty‐seven characters modified from Eberle (1999) and Schoch (1986), and fourteen taxa are presented here with the bremer support values from NONA, without the removal of the outgroups (Figure 14). This initial cladistic analysis has Alveugena

Figure 14. Cimolestid relations tree from cladistic analysis. Single most parsimonious tree found in NONA, with a length of 123, CI of .55, and RI of .68. Bremer supports from NONA added.

remaining as the sister group to the taeniodonts, a hypothesis put forth by Eberle

(1999), and neither of the Procerberus species included is better suited for that position.

This topology holds despite the new evidence from the lower jaw of Alveugena and the addition of the most basal taeniodont, Schowalteria.

I ran a second analysis using the same characters but with added groups outside of the taeniodonts, to test the hypothesis of cimolestid origins of the taeniodonts (i.e.,

that Cimolestes, Procerberus, or a related taxon gave rise to the taeniodonts) versus the

hypothesis of palaeoryctid origins (i.e., that a form such Didelphodus Cope, 1882b;

Acmeodon Matthew & Granger, 1921; Aaptoryctes Gingerich, 1982; or Palaeoryctes

Matthew, 1913 gave rise to the taeniodonts).

In this second analysis, Protictis Matthew, 1937, an early representative of the

Order Carnivora, was chosen as an outgroup because of its distance from the Cimolesta groups in question here. Both McKenna and Bell (1997), a synthetic classification, and

Wible et al. (2007), a cladistic analysis based on many morphological characters, agreed that the carnivorans do not originate from Cimolestes as had been hypothesized by

Lillegraven (1969). However, carnivorans are another group in Ferae that fall outside of

the Cimolesta, and Protictis is well sampled and described, unlike some other early carnivores.

These characters were scored for nineteen taxa, including all known

Taeniodonta, using the literature (Protictis‐ Krause and Gingerich, 1983; Didelphodus–

Van Valen, 1966; Acmeodon‐ Van Valen, 1966 and Wilson, 1985; Aaptoryctes‐ Gingerich,

1982; Palaeoryctes‐ Bloch et al., 2004), the description of the lower dental morphology

of Alveugena (see Appendix A), and observations of taeniodont and other specimens at the AMNH and NMNH.

I ran the analysis with NONA (Goloboff, 1993), using Winclada, which resulted in a single most parsimonious tree with bremer support values calculated. The results of this analysis are summarized in Figure 15. This analysis has Alveugena as a sister group

Figure 15. External relations tree from cladistic analysis. Single most parsimonious tree found in NONA, with a length of 171, CI of .40, and RI of .63. Bremer supports from NONA added. Higher taxa affiliations added.

of the taeniodonts, with Cimolestes falling out as the sister group to that. Also, the

Procerberus species do not fall out as a line leading to Alveugena and the taeniodonts, contrary to Eberle (1999) and the above analysis using fewer taxa.

Other patterns that emerge here include the paraphyly of McKenna and Bell's

(1997) Cimolestidae, which includes not only Cimolestes and Procerberus but also

Didelphodus and Acmeodon, which here lead to the paleoryctids. Although Aaptoryctes and Palaeoryctes are widely thought of as palaeoryctids, the two "cimolestid" taxa falling out with them are commonly disputed as to whether they really are palaeoryctids

or not. However, the paraphyly of the cimolestids is apparent either way in this analysis, and any other inferences about the likelihood of palaeoryctid and cimolestid taxa would require more work. However, it is very possible to point out that given palaeoryctids and cimolestid groups outside of the taeniodonts, taeniodonts do fall closer to the cimolestids and particularly Cimolestes, than to other groups.

Alveugena Sister Grouping

The first part of this study greatly supports Alveugena as the sister group for the taeniodonts, with a Bremer support value of 4. When more outgroups are added, this bremer support value does drop, but it does not drop below the 3 required to be well‐ supported (Wenzel, 2002).

These studies show that Alveugena is most likely the closest sister group to the

Taeniodonta known to this date. Other possible (or once purported) sisters are used in

this study, and Alveugena falls closer than any others. This agrees with Eberle's (1999) placement of Alveugena as the sister group of the taeniodonts. However, the second analysis complicates matters in the cimolestid realm because Cimolestes falls between

Procerberus and Alveugena, whereas it was used as an outgroup in the earlier study

(Eberle, 1999) and the first analysis presented. Because the outgroup in this study

(Protictis) was not any of the taxa whose relation to the taeniodont I was testing, it was possible for Cimolestes to fall closer to the taeniodonts than Procerberus (in Eberle's and the first analysis, Cimolestes was constrained to the base of the tree). Although this study agrees with Eberle (1999) in supporting the hypothesis of cimolestid origins, this study disagrees with Eberle (1999) over which cimolestid (Cimolestes or Procerberus) is more closely related to the taeniodonts.

This new specimen is also from an earlier time period than the original

Alveugena type specimen, which is an upper dentition. Our specimen dates from the very early Puercan, whereas the original is from the Puercan 2 (Eberle, 1999). Since R. C.

Fox and Naylor (2003) claimed that Alveugena's later age would prevent it's placement as the ancestor to the taeniodonts, the earlier age of this new material will help to eliminate this problem or at least diminish ghost lineage length implied by the temporal gap between the oldest taeniodont (Lancian NALMA, or latest part of the Late

Cretaceous) and the appearance of the sister taxon of the taeniodonts (Pu1 interval‐

zone of the Puercan NALMA). Because it is probable that less than a million years actually separate these two genera, Alveugena's or Schowalteria's actual (as opposed to

observed) temporal ranges have overlap, and then Alveugena could be the ancestor to the taeniodonts. There are no autapomorphic characters in Alveugena and absent in the taeniodonts that would preclude Alveugena from being the ancestor to the taeniodonts, and both of these analyses show that it is the taxon most closely related to the

taeniodonts. Thus, it is possible that Alveugena is the ancestor to the taeniodonts.

Taeniodont Monophyly

The analyses presented above solidify the monophyly of Taeniodonta.

Taeniodonta is supported as a whole group, and since there was only one most parsimonious tree, there are no contradictory statements in these parsimony analyses.

The Bremer support for the clade in Figure 14 is a 3, which is a well‐supported grouping.

R. C. Fox and Naylor (2003) speculated that Schowalteria was a stylinodontid, based on characters without a cladistics analysis, and that its early appearance caused a rift in the

monophyly of the group. Here, Schowalteria falls as the sister group to the rest of the taeniodonts, and this agrees with its early arrival in the fossil record. It could be the ancestor to the rest of the group, because, again there are no characters precluding

Schowalteria from being the ancestral taeniodont.

Stratocladistic Analysis

As before, the first results are the same as the previous chapter (Figure 11). The analysis agrees with both of the analyses above in the placement of Alveugena as the sister group to the taeniodonts and in the monophyly of the taeniodonts. The

placement of the Procerberus groups as the sister to Alveugena+Taeniodonta may arise from the use of Cimolestes as the outgroup (as in the differences between the above two analyses). The stratocladistic analysis shows that Cimolestes can be the ancestor, indicated in the figure by horizontal orientation, to the Procerberus and

Alveugena+Taeniodonta, which is a main benefit of stratocladistic analysis. Also, the fact

that the stratigraphic placement of the fossils does not contradict the placement of

Alveugena as the sister group of the taeniodonts is important, especially considering its controversy due to its younger age (R. C. Fox and Naylor, 2003, discussed above).

When the additional outgroups and palaeoryctids are included in a stratocladistic analysis, the same groupings occur. There were many most parsimonious trees, summarized in Figure 16. However, the same placement of Cimolestes as the ancestor to the taeniodonts and Procerberus species occurs, and the other paleoryctids fall out as a sister group to that clade. Similar to the other stratocladistic analysis, the placement of Procerberus formicarum as the sister group or the ancestor of Procerberus grandis both occur, as well as the placement of Protictis as an ancestor or just a sister group, which cause the abundance of most parsimonious trees. The latter issue can be explained by the program trying to force ancestry where none truly occurs (the much younger carnivoran Protictis is not likely the ancestor to Cimolestes). Because Protictis is used as an outgroup, but is later occurring, it is possible that the program tried to force the ancestry, simply because of it's outgroup status. Because it appears in the analysis as both a sister taxa and an ancestor, unlike Cimolestes in the earlier analysis which is

Figure 16. External relations tree from stratocladistic analysis. Summary of most parsimonious trees found in StrataPhy. Procerberus formicarum and Protictis both fall as sister groups or ancestors in this analysis, but are shown here in the sister taxa placement. Horizontal taxa represent ancestral placement.

always an ancestor, it is safe to say that this placement is not as problematic as one might imagine. However, the placement of Cimolestes as an ancestor is definitely supported, because Cimolestes falls out as the ancestor in every parsimonious tree found.

Taeniodont Relations to Cretaceous Stem Eutherians

Because it has been shown that Alveugena is the sister taxon to the taeniodonts and that the cimolestids could be the ancestral group to an Alveugena+Taeniodonta clade, the next step is to try to determine the placement of Taeniodonta among the

Mesozoic eutherian mammal lineages. In the Mesozoic there were many stem eutherian lineages that did not lead to any crown eutherian groups, although some led to taxa in the Cenozoic. Based on its historic affiliation with Cimolestes, it would have been automatically considered a stem eutherian (Wible et al., 2007). A study about the placement of the taeniodonts among eutherian mammals has never been done and will be able to help explain whether the taeniodonts belong with the Mesozoic stem or

Cenozoic crown eutherians.

I did this analysis using a modified .and shortened) version of a published character matrix (Wible et al., 2007). Wible et al. (2007) were looking to determine the placement of many stem and crown eutherians based on morphological characters, and

I adapted their character for this study. Schowalteria and Alveugena were added as the taeniodont and sister group to see where they would fall. Montanalestes was used as

the outgroup because it is an undisputed stem eutherian that is very basal and it is better sampled than other outgroup taxa that were available. The original study had 408 dental, cranial, and postcranial characters (Wible et al., 2007). In this modified analysis, the postcranial characters were removed because the amount of missing data swamped the useful data, leaving 336 characters. The resulting matrix was placed in NONA, and the same parsimony analysis was run as previously. Two most parsimonious trees resulted, and the consensus of which is shown in Figure 17. The only difference between the two most parsimonious trees was the placement of Kennalestes and Asioryctes as either a clade or as a paraphyletic grouping.

Figure 17. External relations among stem Eutheria from cladistic analysis. Strict consensus of two most parsimonious trees found in NONA, each with length 180, CI .85, and RI .75. Groupings have been added to the non‐cimolestid and taeniodont taxa.

The main result of this analysis is that the taeniodonts fall out with Cimolestes.

This analysis places the taeniodonts among stem Eutheria, based on Wible et al.'s (2007) placement of Cimolestes as a stem eutherian. This is the first time a taeniodont has been named as a stem or crown eutherian, and this analysis places them squarely with the

former, along with Alveugena.

Placement within Eutheria

With the taeniodonts placed with other stem Eutherians in the analysis using the modified Wible et al. (2007) matrix, it is important to check this result using early or plesiomorphic representatives of the modern clades of eutherians, to make sure that the group does not fall out among some known later‐appearing group. The same

characters were used as previously and the same analysis was performed. The result was a single most parsimonious tree (Figure 18).

Figure 18. External relations among crown Eutheria from cladistic analysis. Single most parsimonious tree found in NONA, with length 501, CI .65, and RI .49. Higher taxonomic classifications added to the non‐cimolestid and taeniodont taxa.

It is immediately evident that the taeniodonts fall again as a stem taxon. They fall closer to the base than any of these modern groups, though all modern groups are not

represented here. One interesting note is the length of this tree. It may be shocking to some to see a tree length over 500. This results because of the large number of characters in this study, and because the later taxa (particularly the extant taxa) are so

far derived. It is interesting to note that the vast majority of these extra steps occur

after the Cimolestes‐Alveugena‐Schowalteria split from the crown groups in this analysis. The vast majority of the characters holding separate the stem and crown eutherians in this analysis were dental. Several were molar characters, only three were cranial, and one was a premolar character, a diastema present between the first and second lower premolar. The upper molar characters included a strongly reduced or absent ectoflexus, a metacone and paracone that were subequal with bases separated, and weak or absent postmetacrista. The lower molar characters were trigonids that were less than twice or subequal in height and subequal or narrower in width than the

talonid and possessing a protoconid that is subequal to the paraconid and/or metaconid. Finally, the cranial characters were the absence of a condyloid crest, the condylar process existing without a posteriorly directed peduncle and a condyle that lies above the tooth row by more than a molar length. This shows once again that the taeniodonts belong with the stem eutherian mammals and not with some later clade.

This also gives us characters that can be used to distinguish stem from crown eutherians in the future.

44 Chapter 4: Conclusions

The taxon Taeniodonta is a monophyletic group of stem eutherian mammals that

ranged from the Late Cretaceous to the middle Eocene in western North America. The oldest and most primitive taeniodont is from the Late Cretaceous of Canada, and the geologically younger taeniodonts occur in more southern locations. They range all across the western interior, from Montana and North Dakota south to Texas, and have been found in Maryland and South Carolina. Two separate groups evolved from the earliest taeniodont, the Conoryctidae and the Stylinodontidae. The conoryctids are presumed to have evolved in the northern latitudes of North America due to their earliest occurrences there, whereas the stylinodontids first appeared at the southern localities. Both of the groups spread throughout the western interior, and the stylinodontids reached Texas and the east coast.

Although some workers have speculated that the conoryctids may be a paraphyletic group leading to the stylinodontids (e.g., Lucas and Williamson, 1993), all

results here indicate the group (Conoryctella, Conoryctes, and Huerfanodon) excluding

Onychodectes is a solid, monophyletic group separate from the stylinodontids. The conoryctids are smaller and overall are more primitive than the stylinodontids. Although some stylinodontids occurred in the same time periods as the conoryctids (see Figure

45 19), the stylinodontids are much larger, with more robust features and much more

derived dental morphology. The conoryctids retained primitive tribosphenic, insectivorous cheek teeth, whereas in the later stylinodontids cheek teeth become more square and more subject to wear, probably from eating rougher foods, quite possibly using their large claws to dig up roots and tubers. All analyses point to these groupings, and the cladistic and stratocladistic analyses agree with the evolutionary history of taeniodonts (Figure 19).

Figure 19. Known taeniodont stratigraphic distributions with phylogeny. Internal structure reflected found in this study.

46 Patterson (1949), based on the information available to him at the time, inferred that rates of evolution were relatively rapid in the stylinodontids as compared with the conoryctids. Using the concept of adaptive zones, he described the stylinodontids as changing rapidly, particularly between Wortmania and Psittacotherium (Patterson,

1949). Since Patterson's time, new discoveries, such as Schowalteria and Schochia have made it possible to reexamine these evolutionary trends again. Also, with the new phylogenetic information from this study, we can look at the evolutionary patterns. One of the main differences between the conoryctids and stylinodontids in evolutionary pattern is in the earliest divergence from the morphology of the most basal taeniodont

(Figure 20). Because Schowalteria is now known to be the most basal taeniodont, the starting point for evolution of the group has moved since Patterson (1949), who used a hypothetical ancestor in the same adaptive zone as Onychodectes, which was at the

time the earliest known taeniodont. Patterns of character divergence in the taeniodonts are shown in Figure 20, based on canine and first molar size, with probable ancestor– descendent lineages based on the phylogenetic analysis from this study. The early conoryctids exhibited a slower rate of evolution of their molars and canines, whereas

the stylinodontids had a rapid evolution of molars and canines until Psittacotherium, after which stasis occurred, perhaps indicating stabilizing selection. At the same time as the stylinodontids were ending their rapid evolution and moving toward stasis, the conoryctids demonstrated rapid evolution moving between the early conoryctid,

Conoryctella and the later ones, Conoryctes and Huerfanodon. It is possible that once

47 Figure 20. Patterns of character divergence in canine and first molars. Size measured as ln(l*w). Red lines surround the conoryctids of these analyses, blue lines surround the stylinodontids, and green Cretaceous‐ Paleogene (K/P) boundary hand added. Vertical lines indicate evolutionary stasis, perhaps caused by stabilizing selection. Horizontal lines connecting ancestor‐descendent lineages indicate rapid evolutionary change.

48 the stylinodontids had reached their full size and final canine morphology, it left an open niche for larger insectivores while the stylindontids evolved into the digging, tuber‐eaters that began with Psittacotherium.

The origins of the taeniodonts have been debated for many years, commonly because the two groups of taeniodonts seem so different, with the primitive conoryctids

seeming more like early insectivores and carnivores and the stylinodontids are larger, more robust and herbivorous. Placement of taeniodonts within the Cimolesta (McKenna and Bell, 1997), a group of mammals of great variety that are thought to have come from Cimolestes (McKenna, 1975) suggested that taeniodont origins lie within the early

genera of that group, the cimolestids (including Cimolestes). Eberle's (1999) phylogeny

placed Alveugena as the sister taxon to the taeniodonts, with the Procerberus species also closely related. In the analyses contained within, it is evident that Alveugena is the

sister group, but Cimolestes may be more closely related to the taeniodonts than is

Procerberus. Cimolestes and Alveugena are the closest sister groups (Figure 14) to the taeniodonts in these analyses, and quite possibly form a sequence of ancestors and

descendants. No other of the "palaeoryctid" or other cimolestan genera studied fell out

closer, showing that the cimolestids are more closely related to the taeniodonts than other “paleoryctids” or other cimolestans.

49 Systematic Paleontology

The last revision of the taeniodont taxonomy was done more than two decades

ago. With the new information in these studies, including the phylogenetic placement of

all the taeniodonts, it is necessary to rediagnose the clades that have been found and to

include all the known genera that correspond to those clades.

Taeniodonta Cope, 1876

Included genera: Onychodectes Cope, 1888a; Wortmania Hay, 1899; Conoryctes Cope,

1881; Conoryctella Gazin, 1939; Huerfanodon Schoch and Lucas, 1981a; Psittacotherium

Cope, 1882a; Ectoganus Cope, 1874; Stylinodon Marsh, 1874; Schochia Lucas and

Williamson, 1993; and Schowalteria R. C. Fox and Naylor, 2003.

Distribution: Late Cretaceous to Uintan (middle Eocene) of western North America,

Tiffanian (middle to late Paleocene) and Uintan of Texas, and Tiffanian of South

Carolina.

Revised Diagnosis: Animals larger than the cimolestids that differ from Cimolestes and

Alveugena in the following ways: Canines moderate to large, stout, recurved; P1/p1

single rooted, P2/p2 double rooted, P3 triangular, premolariform p3, P4 with well‐ developed paracone and protocone, lacking a metacone, with cuspate stylar shelf, and

p4 nonmolariform with a well developed talonid heel, more so than the cimolestids;

upper molars molariform with small, lingually placed protocone and conules, paracone

50 and metacone moderate and placed far labial, small stylar shelf, no pre‐ or post‐cingula,

ectocingula present and well developed; lower molars with variable relative

trigonid/talonid widths, trigonid/talonid heights generally subequal and where different

never as different as with the cimolestids, protoconid and metaconids subequal with

smaller paraconids placed more labial than the metaconid, lacking the accessory talonid

notch cusp of Cimolestes and Alveugena, and in m3 talonid not expanded; hypsodonty

found in all cheek teeth, with early animals showing more crown hypsodonty and later

showing more root hypsodonty to hypselodonty; wear pattern generally over entire

surface of cheek teeth and often deep enough to expose pulp cavity.

Comments on Diagnosis: This diagnosis has been revised from Schoch's (1986) diagnosis.

Premolar, canine, and wear characters were added, and also lower molar paraconid characters. Characters removed include subequal talonid and trigonid on the lower molars and decreasing molar size posteriorly.

Discussion: Taeniodonta is here referred to as a monophyletic clade as demonstrated above. The new discovery of Schowalteria has modified this diagnosis the most, as it is

the earliest and most basal taeniodont, whereas Onychodectes had been most basal at

the time of the last diagnosis (Schoch, 1986). Although the monophyly of this group has

been debated, these are the first such parsimony analyses of the entire group; and,

therefore, show that this group is monophyletic.

51 Taeniodonta Cope, 1876

Conoryctidae Wortman, 1896

Type genus: Conoryctes Cope, 1881.

Included genera: Conoryctella Gazin, 1939; Conoryctes Cope, 1881; and Huerfanodon

Schoch and Lucas, 1981a.

Distribution: Torrejonian (middle Paleocene) of western North America.

Revised diagnosis: Smaller than the stylinodontids, probably insectivorous taeniodonts with incipient metacone on P4 and loss of all stylar shelf and associated cusps, p4 with

small talonid heel; upper molars with subequal metacones and paracones, reduced

stylar shelf and small mesostyle.

Comments: Diagnosis revised from Schoch (1986) with addition of weak or absent stylar

shelf on molars and P4 loss of stylar shelf and all smaller cusps and the removal of

crown hypsodonty and triangular upper premolars.

Taeniodonta Cope, 1876

Stylinodontidae Marsh, 1875

Type genus: Stylinodon Marsh, 1874

Included genera: Wortmania Hay, 1899; Schochia Lucas & Williamson, 1993;

Psittacotherium Cope 1882a; Ectoganus Cope, 1874; and Stylinodon Marsh, 1874.

Distribution: Puercan (early Paleocene) to Uintan (middle Eocene) of western North

America, Tiffanian (middle to late Paleocene) and Uintan of Texas, and Tiffanian of

South Carolina.

Revised diagnosis: Larger bodied taeniodonts with large canines; lower premolars oblique, multi‐cusped and more complex than all other taeniodonts; upper molar ectocingula slight to absent, stylar shelf weak or absent; root hypsodonty including deep rooted cheek teeth and canines, quickly developed into canine hypselodonty and finally cheek teeth hypselodonty as well; short, deep skulls; large claws on manus and robust limb bones.

Comments: Diagnosis revised from Schoch (1986) with addition of root hypsodonty, multicusped premolars, and upper molar ectocingulum characters.

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Appendix A: Description of the lower jaw of Alveugena

SYSTEMATIC PALEONTOLOGY Order CIMOLESTA McKenna, 1975 Suborder DIDELPHODONTA McKenna, 1975 Family CIMOLESTIDAE (Winge, 1917) Simpson, 1931 Genus ALVEUGENA Eberle, 1999

Type and only known species‐ Alveugena carbonensis.

Revised diagnosis

Upper dentition diagnosed by Eberle (1999) as "Includes the largest documented

cimolestid; upper dentition morphologically intermediate between the cimolestid

didelphodont Procerberus and the taeniodont Onychodectes; dental formula for upper dentition is I2, C, P4, and M3; on P3–4, paracone is largest cusp, occupying over half the transverse width of tooth, and metacone is incipient or absent; P3–4 with weak para‐ and metastyles and narrow, discontinuous stylar cingula; pronounced wear pattern over most of crown on P3–M3, as in taeniodonts; weak molar ectoflexus; molar conules labial, molars with antero‐ and posterolingual cingula; M1 with continuous lingual cingulum; M2 shorter, but more transverse, than MI; M3 smaller than MI–2; upper molars more transverse than Onychodectes"; dental formula for lower dentition is i? c1

p4 m3; a moderate, slightly recurved and ovoid canine, in a trend from smaller, less

recurved Cimolestes to larger, more recurved Schowalteria; p1 single rooted and round;

p2–4 double‐rooted, nonmolariform, talonids ranging from very small to well developed respectively, p4 talonid not as well developed as in Schowalteria, talonids lack the extra cusp of Cimolestes; m1–3 double‐rooted with precingulid, m1–2 paraconid smallest cusp slightly lingual of center with lower base, metaconid larger, protoconid largest, talonid

basin with accessory cusp in talonid notch as in Cimolestes, but not between the

hypoconid and hypoconulid as in Procerberus grandis.

ALVEUGENA CARBONENSIS Eberle, 1999

Diagnosis

Same as genus.

Description

The specimen contains c1–m3 and some dentary bone. The p1 is missing. The anterior portion of the p4 protoconid has been broken, and the posterior portion of m3 is missing.

The canine of this specimen is long, deep rooted, slightly recurved, and ovoid in

cross‐section. The long root lifts the crown above the rest of the jaw line (though this may be from breakage and repair). The crown itself is slightly larger than those of the molars, and the enamel reaches farther down the tooth on the buccal side. The canine

crown is smaller than in Schowalteria clemensi R. C. Fox and Naylor, 2003, where the

canine crown almost doubles the height of the molars. Alveugena's canine is also slightly higher and thicker than that of Cimolestes Marsh, 1889. The recurve of this canine falls

Figure 21. Alveugena lower jaw in buccal, lingual, and occlusal view. Scale bar is 1 cm.

between that of Cimolestes and Schowalteria, along a trend moving toward more recurved through this series.

This animal had four premolars. The alveolus of p1 shows that it was single rooted, small, and circular in cross‐section, similar to Cimolestes cerberoides Lillegraven,

1969. Schowalteria's p1 is more complex, although still single rooted, but not circular.

The rest of the premolars are all double‐rooted, as with Cimolestes and Schowalteria.

The p2 consists of a large, slightly recurved protoconid and a very small talonid heel, similar but less robust than Schowalteria with a smaller talonid heel and lacking an

anterior ridge. There is an extra accessory cusp on the talonids of Cimolestes p2‐4, which would be more complex than that of Alveugena. However, Procerberus lacks these extra cusps on p3–4, so it is probable that they are also absent on the yet undescribed p1–2 of Procerberus (Lillegraven, 1969). The p3 also has a large protoconid

and small talonid heel, again similar to Schowalteria, except for robustness and an anterior ridge. It is difficult to determine the height of the protoconid due to wear, but the width is also consistent with Cimolestes magnus. The talonid of Alveugena is better

developed. The p3 may have once had one cusp on the talonid as in Procerberus, but the surface of the talonid has been removed by wear. There are small diastemata between p1–3 as in Schowalteria. The nonmolariform p4 consists of a larger protoconid, broken anteriorly, and a talonid heel that is more well developed than p3 but still small.

The p4 in Procerberus is submolariform so it is vastly different from the p4 of Alveugena.

This talonid falls between the development of Schowalteria, which is very well developed, and Cimolestes magnus, which is only moderately sized. Schowalteria also

contains a precingulid that Alveugena lacks.

The m1–3 are double‐rooted with a low precingulid near the bottom of the

protoconid. The m2 is very similar to the m1 except much less worn, and m3 exists only

as a protoconid. Though the trigonid of m1 is worn flat, it is possible to determine that

the cusps decrease in size from protoconid to metaconid to paraconid, and the m2 has a

trigonid is approximately twice the height of the talonid, with a tall protoconid, medium

metaconid, and smaller paraconid. This character state differs from that of Procerberus

grandis and Schowalteria, which have subequal metaconids and paraconids, but it is consistent with that of Cimolestes magnus. The paraconid is again slightly lingual to center and farther down on the anterior of the tooth, as in Cimolestes magnus and

Procerberus grandis. Although the talonid is worn in both teeth, it is still possible to

determine that the hypoconid is large (approximately the same size as the paraconid), and the hypoconulid and entoconid are smaller and subequal in size. The m1–2 have a

slight upward protrusion near the bottom of the entoconid, which is also in Cimolestes

magnus and Procerberus grandis, although the latter has another accessory cusp

between the hypoconulid and hypoconid that is not present here.

The dentary is only visible from below p1 to halfway below m3, so the only

distinct characteristics are mental foramina. There are two present, one below the

anterior portion of p1, similar to Onychodectes, and the other below the center of p4,

similar to Cimolestes. Schowalteria also has two mental foramina, but they are closer together, located between p1 and p2 and then between p3 and p4.

Comparison to UW 26497 and 26269

This jaw has a striking resemblance to UW 26497 and 26269, which was thought

to probably occlude with the uppers of Alveugena (Eberle and Lillegraven, 1998). Our specimen is slightly smaller in overall size, including tooth sizes, but not eough to rule our their belonging to a single species. The tall protoconid on our molars is similar to

those on the UW specimens. The main difference between them is that our specimen is

much less damaged. The UW specimen seems to have had much more damage between death and burial.

Wear Descriptions

The only wear present on the canine is on the tip, which is angled 45 degrees down the posterior side, similar to Schowalteria. On the p2, there is wear on the

posterior side of the protoconid, which is shared with Schowalteria, although the latter

has several other wear facets not in Alveugena. The protoconid of the p3 is worn almost

flat, although there is a slight rise anteriorly at approximately 25 degrees. This main

wear continues to the anterior side of the talonid, creating a single wear facet. A similar wear pattern as p3 exists on p4, except to a lesser degree. Wear on the talonid obscures any detail but does not make a continual wear facet, as in Schowalteria, because there is enamel still visible on the vertical posterior face of the protoconid. Schowalteria has more wear facets than Alveugena, which generally include both anterior and posterior

sloping facets on both sides of the premolar protoconids. This type of wear is not present in Alveugena, which only has posterior sloping wear.

The trigonid of m1 is almost completely worn flat, except for a patch of enamel

between the metaconid and paraconid. The talonid is worn flat except for the talonid notch. The paraconid of m2 is worn anteriorly but not to flatness. The talonid has less

wear. The basin is worn from the interior sides of the conids inward, but there is enough

left to show that no cristids were present on the talonid. The hypoconulid also has some

posterior wear, which leaves it almost flat. The trigonid wear patterns are similar to those of Schowalteria in having both an anterior wear that slopes anteriorly and a posterior wear that slopes posteriorly. This would account for the shape of the flat surface of m1, with the anterior portion sloping anterior and the poster section more

flat, but sloping slightly posterior. However, the talonid wear differs from Schowalteria

because it is separated from the posterior slope. This second wear facet more closely

resembles that of Cimolestes magnus and Procerberus grandis, as a cut from the talonid

notch to between the hypoconulid and hypoconid, but wider than in those species.

Appendix B: Characters and Matrices

Internal/External Analyses Additive characters, where evolutionary steps have to move through intervening character states, marked with an asterix (*). Characters coded from Schoch (1986) for all taxa, and those from Eberle (1999) coded for all taxa excluding Cimolestes, Procerberus formicarum, Procerberus

grandis, Alveugena, and Onychodectes, which appeared in the original analysis.

[1] Upper molar protocone size (Schoch, 1986) * 0= small 1= moderate 2= large [2] Upper molar conule size (Schoch, 1986) 0= small 1= large 2= absent [3] Placement of conules on upper molars (modified from Eberle, 1999) 0= labial 1= lingual [4] Upper molar paracone size (Schoch, 1986) * 0= small 1= moderate 2= large [5] Stylar margin on upper molars (Eberle, 1999) * 0= large, inflated stylar lobes; forward‐projecting, parastylar lobe extends labially beyond, and often "hooks" around, metastylar lobe on tooth directly in front of it; stylar lobes appear largest on M2. 1= para‐ and metastylar lobes relatively smaller than those of Cimolestes and Procerberus formicarum, hut still relatively inflated; parastylar lobe is not hook like, does not project anteriorly, and projects only slightly more labially (if at all) beyond metastylar lobe of tooth directly anterior to it. 2= small, weak, uninflated stylar lobes; stylar shelves very narrow to absent; M2 stylar lobes subequal in size to those of other molars. [6] Upper molar metacone size (Schoch, 1986) * 0= small 1= moderate 2= large 66

[7] Lingual cingula on upper molars (Eberle, 1999) 0= present 1= absent [8] Lower molar trigonid height (modified from Eberle, 1999) * 0= trigonid noticeably higher than talonid 1= trigonid only slightly higher than talonid 2= trigonid equal in height to talonid [9] Lower first molar trigonid width (modified from Eberle, 1999) 0= trigonid wider than talonid 1= trigonid equal in width to talonid 2= width variable in genus [10] Lower second molar trigonid width (modified from Eberle, 1999) 0= trigonid wider than talonid 1= trigonid equal in width to talonid 2= width variable in genus [11] Lower third molar trigonid width (modified from Eberle, 1999) 0= trigonid wider than talonid 1= trigonid equal in width to talonid 2= width variable in genus [12] Paraconids versus metaconids (Schoch, 1986) 0= not equal 1= subequal [13] Molar hypsodonty (modified from Eberle, 1999) * 0= absent 1= present 2= hypselodonty [14] P3 morphology (modified from Eberle, 1999) * 0= triangular 1= submolariform 2= molariform [15] P4 morphology (modified from Eberle, 1999) * 0= triangular 1= submolariform 2= molariform [16] p4 talonid heel (Schoch, 1986) * 0= small 1= moderate 2= well=developed [17] P4 parastyle (modified from Eberle, 1999) * 0= absent 1= incipient 2= small 3= well=developed [18] P4 stylocone (modified from Eberle, 1999) * 0= absent

1= incipient 2= small 3= well=developed [19] P4 metastyle (modified from Eberle, 1999) * 0= absent 1= incipient 2= small 3= well=developed [20] Paraconids (Schoch, 1986) * 0= small 1= moderate 2= large [21] P1 (Schoch, 1986) 0= present 1= absent [22] p4 morphology (Schoch, 1986) * 0= nonmolariform 1= submolariform 2= molariform [23] M1 and M2 mesostyle (Schoch, 1986) 0= absent 1= small 2= moderate 3= well=developed [24] Premolars set obliquely (Schoch, 1986) 0= no 1= yes [25] Lower canines (Schoch, 1986) * 0= small 1= moderate 2= large [26] Upper incisor number (Schoch, 1986) * 0= 3 1= 2 2= 1 [27] Lower incisor number (Schoch, 1986) * 0= 3 1= 2 2= 1 [28] p3 morphology (Schoch, 1986) * 0= nonmolariform 1= submolariform 2= molariform [29] Lower canine root (Schoch, 1986) * 0= shallow

1= deep 2= tending towards rootlessness 3= rootless [30] Bilophodont molars (Schoch, 1986) 0= absent 1= present [31] Transverseness of upper molars (Eberle, 1999) * Relative Transverse Width (RTW)= Maximum Transverese Width/A‐P length 0= transverse (RTW of M1 > 1.30) 1= less transverse (RTW of M1 > 1.20) 2= weakly to nontransverse (RTW of M1 < 1.20) [32] Development of molar ectoflexus (Eberle, 1999) * 0= pronounced and deep, particularly on M2 1= small, shallow indentation 2= virtually absent, resulting in a relatively straight stylar shelf [33] Size of M2 relative to M1 (Eberle, 1999) * 0= M2 noticeably larger than M1 1= M2 shorter but more transverse than M1 2= M2 subequal in size to, or slightly smaller than, M1 [34] Upper molar lingual length (modified from Eberle, 1999) 0= not pronounced 1= pronounced [35] Metacrista on upper molars (modified from Eberle, 1999) * 0= well=developed 1= reduced 2= absent [36] Lower molar cingulids (Eberle, 1999) 0= present 1= absent [37] Wear pattern on upper molars and premolars (Eberle, 1999) 0= not pronounced over entire occlusal surface 1= pronounced over entire occlusal surface

Matrix for internal structure of Taeniodonta‐ Cladistic Analysis Cimolestes ?100000000000001333000002?00000000000 Procerberus formicarum 010000000000022?333?0000???0001000100 Procerberus grandis 01001001000?0222333200001??0001000101 Alveugena 0?01110000?000002320000011?0001111101 Schowalteria ????1?11100?1002232000?021?1000001?11 Onychodectes 0101211122011011121200001000002221111 Conoryctella 2202221122201011232201102000201121111 Conoryctes 2202221120001121232011202110100221111 Huerfanodon 02122211200111?2???111302??0102221011 Schochia 1102201?????102?010???01??????02211?1 Wortmania 10-2201000?11002111100012220010211111 69

Psittacotherium 2212201220001112010001012221210221111 Ectoganus 2212221220211222010002012222312221211 Stylinodon ????2?12111?2222010?02?12122312221?11

Matrix for cimolestid v. palaeoryctid placement of Taeniodonta‐ Cladistic Analysis Protictis 1102000000010111303101101000100111110 Cimolestes ?10000000000000133300000???0000000000 Acmeodon 0202101000000012???111101??0?00010000 Aaptoryctes 2110001000000011000011000?00000010100 Palaeoryctes 22?20010000001113300?0001?00000010000 Didelphodus 0202000000000011202101001001000000000 Procerberus formicarum 010000000000022?333?0000???0001000100 Procerberus grandis 01001001000?0222333200001??0001000101 Alveugena 0?01110000?000002320000011?0001111101 Schowalteria ????0?11100?1002232000?021?1000001?11 Onychodectes 0101211122011011121200001000002221111 Conoryctella 2202221122201011232201102000201121111 Conoryctes 2202221120001121232011202110100221111 Huerfanodon 02122211200111?2???111302??0102221011 Schochia 1102201?????102?010???01??????02211?1 Wortmania 10-2201000?11002111100012220010211111 Psittacotherium 2212201220001112010001012221210221111 Ectoganus 2212221220211222010002012222312221211 Stylinodon ????2?12111?2222010?02?12122312221?11

Matrix for internal structure of Taeniodonta‐ Stratocladistic Analysis Cimolestes ?100000000000001333000002?000000000000 Procerberus formicarum 010000000000022?333?0000???00010001002 Procerberus grandis 01001001000?0222333200001??00010001012 Alveugena 0?01110000?000002320000011?00011111012 Schowalteria ????1?11100?1002232000?021?1000001?111 Onychodectes 01012111220110111212000010000022211113 Conoryctella 22022211222010112322011020002011211114 Conoryctes 22022211200011212320112021101002211115 Huerfanodon 02122211200111?2???111302??01022210115 Schochia 1102201?????102?010???01??????02211?13 Wortmania 10-2201000?110021111000122200102111113 Psittacotherium 22122012200011120100010122212102211116 Ectoganus 22122212202112220100020122223122212117 Stylinodon ????2?12111?2222010?02?12122312221?118

Matrix for cimolestid v. palaeoryctid placement of Taeniodonta‐ Stratocladistic Analysis Protictis 11020000000101113031011010001001111105 Cimolestes ?10000000000000133300000???00000000000

Acmeodon 0202101000000012???111101??0?000100005 Aaptoryctes 2110001000000011000011000?000000101005 Palaeoryctes 22?20010000001113300?0001?000000100006 Didelphodus 02020000000000112021010010010000000008 Procerberus formicarum 010000000000022?333?0000???00010001002 Procerberus grandis 01001001000?0222333200001??00010001012 Alveugena 0?01110000?000002320000011?00011111012 Schowalteria ????0?11100?1002232000?021?1000001?111 Onychodectes 01012111220110111212000010000022211113 Conoryctella 22022211222010112322011020002011211114 Conoryctes 22022211200011212320112021101002211115 Huerfanodon 02122211200111?2???111302??01022210115 Schochia 1102201?????102?010???01??????02211?13 Wortmania 10-2201000?110021111000122200102111113 Psittacotherium 22122012200011120100010122212102211116 Ectoganus 22122212202112220100020122223122212117 Stylinodon ????2?12111?2222010?02?12122312221?118

Ancient/Modern Analyses Dental and cranial characters used from Wible et al. (2007). The original study had 408 dental, cranial, and postcranial characters. In this modified analysis, the postcranial characters were removed because the amount of missing data overwhelmed the useful data, leaving 336 characters. Characters coded here for Alveugena and Schowalteria, all

others coded by Wible et al. (2007).

1. Teeth 0= present 1= absent 2. Teeth 0= differentiated into morphological types (is, cs, ps, and ms) with enamel 1= simple peg‐like without enamel 3. Number of postcanine tooth loci 0= eight or more 1= seven 2= six 3= five or less 4. Upper diastema 0= small, between incisors and canine 1= small, between canine and premolars 2= enlarged 3= absent (3). 5. Lower diastema behind incisors 0= absent or small 1= enlarged Dentition – Incisors 6. Incisor shape 0= root and crown are straight and continuous in length 1= a continuous curve 7. Number of upper incisors 0= five 1= four 2= three 3= two 4= one 5= none 8. Number of lower incisors 0= four

1= three 2= two, anterior positions 3= one 4= none or posterior position(s) only 9. Anteriormost upper incisor alveoli 0= approximating 1= separated by a broad gap (1). 10. Anteriormost upper incisor size 0= small, subequal to subsequent 1= enlarged 2= smaller than subsequent 11. Anteriormost upper incisor shape 0= conical 1= mediolaterally compressed 2= anteroposteriorly compressed 3= cuspate (one major and one minor) 4= spatulate 12. Anteriormost upper incisor growth 0= rooted 1= open rooted, in premaxilla only 2= open rooted, extending into maxilla 13. Anteriormost upper incisor enamel 0= surrounds tooth 1= discontinous posteriorly 14. Ultimate upper incisor 0= in premaxilla 1= between maxilla and premaxilla 2= in maxilla 15. Anteriormost lower incisor size 0= small, subequal to subsequent incisors 1= greatly enlarged 2= or tiny, smaller than subsequent 16. Anteriormost lower incisor shape 0= conical 1= mediolaterally compressed 2 =anteroposteriorly compressed 3= cuspate (one major and one minor) 4= spatulate 17. Procumbent anteriormost lower incisor 0= absent 1= present 18. Anteriormost lower incisor root 0= closed 1= open 19. Anteriormost lower incisor root length

0= not extended posteriorly below p1 1= extending posteriorly below p1 2= extending posteriorly below penultimate or ultimate premolar 3= extending posteriorly below molars 20. Anteriormost lower incisor enamel 0= covers the whole incisor 1= discontinous posteriorly 21. Procumbent posterior lower incisor(s) 0= absent 1= present 22. Staggered lower incisor 0= absent 1= present Dentition – Canine 23. Upper canine 0= present, large 1= present, small 2= absent 24. Number of upper canine roots 0= two 1= one 25. Lower canine 0= present, large 1= present, small 2= absent 26. Number of lower canine roots 0= two 1= one 27. Procumbent lower canine 0= absent 1= present 28. Deciduous canine 0= present 1= absent Dentition – Premolars 29. Number of premolars 0= five or more 1= four 2= three 3= two 30. Replacement of dP1/dp1 and dP2/dp2 0= present 1= absent 31. Tall, trenchant premolar 0= ultimate premolar

1= penultimate premolar 2= absent 32. Procumbent first upper premolar 0= absent 1= present 33. First upper premolar roots 0= two 1= one 2= three 34. Diastema posterior to first upper premolar 0= absent 1= present 35. Third upper premolar roots (only scored for taxa with five upper premolars) 0= two 1= one 36. Penultimate upper premolar protocone 0= absent 1= small lingual bulge 2= with an enlarged basin 37. Penultimate upper premolar metacone 0= absent 1= swelling 2= large 38. Penultimate upper premolar parastylar lobe 0= absent or small 1= well developed 39. Penultimate upper premolar roots 0= two 1= three 2= one 3= four 40. Ultimate upper premolar protocone 0= absent or narrow cingulum 1= shorter than paracone 2= approaches paracone in height 41. Ultimate upper premolar metacone 0= absent 1= swelling 2= large 42. Ultimate upper premolar para‐ and metastylar lobes 0= absent or insignificant 1= subequal 2= parastylar lobe larger 3= metastylar lobe larger 43. Ultimate upper premolar precingulum

0= absent 1= present 44. Ultimate upper premolar postcingulum 0= absent 1= present, lower than protocone 2= present, level with protocone 45. Ultimate upper premolar conules 0= weak or absent 1= prominent 46. Ultimate upper premolar size (occlusal surface) relative to first upper molar 0= smaller or subequal 1= larger 47. First lower premolar orientation 0= in line with jaw axis 1= oblique 48. First lower premolar roots 0= two 1= one 49. Diastema separating first and second lower premolars 0= absent (gap less than one tooth root for whichever is smaller of adjacent teeth) 1= present, subequal to one tooth‐root diameter or more 50. Third lower premolar size to second (only scored for taxa with five lower premolars) 0= longer 1= shorter 51. Third lower premolar roots (only scored for taxa with five lower premolars) 0= two 1= one 52. Penultimate lower premolar paraconid 0= indistinctive or absent 1= present and distinctive 53. Penultimate lower premolar metaconids 0= absent 1= swelling 2= separate from protoconid 54. Penultimate lower premolar talonid cusps 0= one 1= two 2= three 55. Ultimate lower premolar paraconid 0= indistinctive or absent 1= distinctive but low 2= distinctive and high 56. Ultimate lower premolar metaconids 0= absent

1= swelling 2= large 57. Ultimate lower premolar talonid 0= narrower than anterior portion of crown 1= as wide as anterior portion of crown 58. Ultimate lower premolar talonid cusps 0= one 1= two 2= three 59. Length of ultimate lower premolar to penultimate 0= longer 1= equal to or less 60. Ultimate lower premolar anterolingual cingulid 0= absent 1= present Dentition – Molars 61. Number of molars 0= four or more 1= three 2= two 62. Size of molar series 0= subequal 1= posterior increase 2= posterior decrease 63. Molar cusp form 0= sharp, gracile 1= inflated, robust 2= crest‐like 64. Upper molar shape 0= as long as wide, or longer 1= wider than long (length more than 75% but less than 99% of width) 2= much wider than long (length less than 75% of width) 65. Size (labiolingual width) of upper molar labial stylar shelf at maximum 0= 50% or more of total transverse width 1= less than 50% but more than 25% 2= less than 25% 2= absent 66. Labial extent of parastylar and metastylar lobes 0= parastylar lobe more labial 1= subequal 2= metastylar lobe more labial 3= lobes absent 67. M1 parastylar lobe relative to paracone 0= parastylar lobe is anterolabial to paracone 1= parastylar lobe is anterior to paracone

68. Length of parastylar lobe (measured to stylocone or stylocone position) relative to total length on penultimate molar 0= more than 30% 1= less than 30% but more than 20% 2= 20% or less 69. Preparastyle 0= absent 1= present 70. Stylar cusp A 0= subequal to larger than B 1= distinct, but smaller than B 2= vestigial to absent 71. Stylar cusp B relative to paracone 0= smaller but distinctive 1= vestigial to absent 2= subequal 72. Stylar cusp C, mesostyle 0= absent 1= present 73. Stylar cusp D 0= absent 1= smaller or subequal to B 2= larger than B 74. Stylar cusp E 0= directly lingual to D or D‐position 1= distal to D 2= small to indistinct 75. Preparacingulum 0= absent 1= interrupted between stylar margin and paraconule or paraconule position 2= continuous 76. Deep ectoflexus 0= present only on penultimate molar 1= on penultimate and preceding molars 2= strongly reduced or absent 77. Metacone size relative to paracone 0= noticeably smaller 1= slightly smaller 2= subequal or larger 3= absent or merged with paracone. 78. Metacone position relative to paracone 0= labial 1= approximately at same level 2= lingual 79. Metacone and paracone bases

0= adjoined 1= separated 80. Preparacrista 0= strong, from side of paracone to stylocone 1= weak, from base of paracone, or absent 81. Cuspate preparacrista 0= present 1= absent 82. Centrocrista 0= straight 1= V‐shaped 2= absent 83. Postmetacrista 0= prominent, from side of metacone to metastyle 1= salient 2= weak, from base of metacone, or absent 84. Cuspate postmetacrista 0= present 1= absent 85. Preprotocrista 0= does not 1= does extend labially passed base of paracone (double rank prevallum/postvallid shearing) 2= absent 86. Postprotocrista 0= extends to mid‐lingual surface of metacone 1= extends distal to metacone 2= absent 87. Development of postvallum shear 0= present but only by the first rank: postmetacrista 1= present, with the addition of a second rank (postprotocrista below postmetacrista) but the second rank does not reach labially below the base of the metacone 2= present, with second rank extending to metastylar lobe: metacingulum 3= absent 88. Paraconule 0= weak or absent 1= prominent, closer to protocone 2= prominent, midway or closer to paracone 89. Metaconule 0= weak or absent 1= prominent, closer to protocone 2= prominent, midway or closer to metacone 90. Internal conular cristae 0= indistinct 1= distinctive and wing‐like

91. Anteroposterior width of conular region (with or without conules) (Luo and Wible, 2005:104) 0= narrow (anteroposterior distance less than 0.30 of total tooth length) 1= moderate development (distance = 0.31–0.50 of total tooth length) 2= wide (distance greater than 0.51 of total tooth length) 92. Protocone 0= lacking 1= small, without trigon basin 2= with distinct trigon basin 93. Protocone antero‐posterior expansion 0= none, subequal to paracone 1= expanded, larger than paracone 94. Protocone procumbency 0= absent 1= present 95. Degree of labial shift of protocone (distance from protocone apex to lingual border vs. total tooth width, in %) 0= no labial shift (10%‐20%) 1= moderate labial shift (21%‐30%) 2= substantial labial shift (≥ 31%) 96. Protocone height 0= low 1= tall, approaching paracone and metacone 2= subequal 97. Precingulum 0= absent or weak 1= present 2= present, reaching labially passed the paraconule or paraconule position 98. Postcingulum 0= absent or weak 1= present, lingual to metaconule or metaconule position 2= present, reaching labially passed metaconule or metaconule position 3= present, extending to labial margin 99. Hypocone on postcingulum 0= absent 1= present, lower than protocone 2= present, subequal to protocone 100. Pre‐ and postcingulum 0= separated 1= continuous lingually 101. Number of penultimate roots 0= three 1= four 2= more 102. Number of roots on ultimate molar 0= three

1= two 2= one 4= four or more 103. Lingual root position on upper molars 0= supporting paracone 1= supporting trigon 104. Ultimate upper molar width relative to penultimate molar 0= subequal 1= smaller 105. Metastylar lobe on ultimate molar 0= absent 1= present 106. Paraconid 0= present 1= absent 107. Paraconid height relative to metaconid 0= shorter 1= subequal 2= taller 108. Paraconid on lingual margin 0= absent 1= present 109. Mesiolingual vertical crest of paraconid 0= rounded 1= keeled 110. Paracristid 0= notched 1= continuous curve without notch 111. Trigonid configuration 0= open, with paraconids anteromedial, paracristid‐protocristid angle more than 50° 1= more acute, with paraconid more posteriorly placed, paracristid‐protocristid angle between 36 and 49° 2‐ anteroposteriorly compressed, paracristid‐protocristid angle 35° or less 112. Protoconid height 0= tallest cusp on trigonid 1= subequal to para‐ and/or metaconids 2= smaller than para‐ and/or metaconid 113. Protocristid orientation 0= oblique 1= transverse 114. Anterior and labial (mesio‐buccal) cingular cuspule (f) 0= present 1= present with a distinct cingular shelf posteroventrally directed from it 2= present with shelf continuing along buccal border 3= absent

115. Talonid 0= small heel 1= multicusped basin 116. Cristid obliqua 0= incomplete, with distal metacristid present 1= complete, attaching lingual to notch in protocristid 2= complete, attaching labial to notch in protocristid 3= complete, attaching below middle posterior of protoconid 4= complete, labially placed 117. Trigonid height relative to talonid height 0= twice or more 1= less than twice 3= subequal 118. Anteroposterior shortening at base of trigonid relative to talonid 0= trigonid long (more than 75% of tooth length) 1= some shortening (50‐75% of tooth length) 2= anteroposterior compression of trigonid (less than 50% of tooth length) 119. Talonid width relative to trigonid 0= very narrow, subequal to base of metaconids 1= narrower 2= subequal to wider 120. Hypoconulid 0= absent 1= in posteromedial position (near the mid‐point of transverse talonid width 2= lingually placed with slight approximation to entoconid 3= close approximation to entoconid 121. Hypoconulid of ultimate molar 0= short and erect 1= tall and sharply recurved 2= posteriorly procumbent 3= absent 122. Entoconid 0= absent 1= smaller than 2= subequal to larger than hypoconid and/or hypoconulid 123. Postcristid (between entoconid and hypoconulid) taller than hypoconulid and nearly transverse 0= absent 1= present 124. Mesoconid 0= absent 1= present 125. Hypolophid 0= absent 1= present

126. Labial postcingulid 0= absent 1= present 127. Ultimate lower molar size relative to penultimate lower molar 0= subequal or larger 1= smaller Mandible 128. Number of mental foramina 0= two or more 1= one 129. Anteriormost mental foramen 0= below incisors (or anteriormost mandible) 1= below p1 2= below p2 3= more posterior 130. Posteriormost mental foramen 0= in canine and anterior premolar (premolariform) region (in saddle behind canine eminence of mandible) 1= below penultimate premolar (under anterior end of functional postcanine row) 2= below ultimate premolar 3= at ultimate premolar and first molar junction or more posterior (3). 131. Depth of mandibular body 0= slender and long 1= deep and short 132. Space between ultimate molar and coronoid process 0= absent 1= present 133. Coronoid process height 0= higher than condyle 1= even with condyle 134. Coronoid process width 0= broad, roughly two molar lengths 1= narrow, subequal to or less than one molar length 135. Tilting of coronoid process (measured as angle between anterior border of coronoid process and horizontal alveolar line of all molars) 0= strongly reclined and angle obtuse (≥150o) 1= less reclined (135o‐ 145o) 2= less than vertical (110o‐125o) 3= near vertical (95o to 105o) 4= tilted anteriorly 136. Coronoid crest 0= absent or weakly developed 1= present and laterally flaring 137. Ventral border of masseteric fossa 0= absent

1= present as a low and broad crest (more than half the height of mandibular ramus) 2= present as a well‐defined and thin crest (less than half the height) 138. Anteroventral extension of masseteric fossa 0= absent 1= extending anteriorly onto mandibular body 139. Labial mandibular foramen 0= absent 1= present 140. Condyloid crest 0= absent 1= present 141. Posterior shelf of masseteric fossa 0= absent 1= present 142. Angular process 0= process on posterior aspect of mandibular ramus 1= shelf along ventral border of mandibular ramus 143. Angular process orientation 0= posteriorly directed 1= medially inflected 2= posteroventrally directed 3= posterodorsally directed 144. Angular process length 0= less than mandibular ramus length 1= equal or greater than mandibular ramus length 145. Angular process shape 0= tapers, base wider than tip 1=rounded, base as wide as tip 146. Angular process vertical position 0= at posteroventral border of mandible 1= posterodorsal, at or near the alveolar border 147. Root of angular process relative to condylar process 0= level with or posterior to 1= anterior to 148. Condylar process 0= with posteriorly directed peduncle 1= not 149. Condyle shape 0= ovoid 1= cylindrical 2=anteroposteriorly elongate 150. Condyle position relative to tooth row 0= at about same level 1= slightly above 2= above by more than molar length

151. Mandibular symphysis shape 0= tapered 1= deep 152. Mandibular symphysis posterior extent 0= p1 or more anterior 1= p2 2= p3 or more posterior 153. Mandibular symphysis 0= mobile 1= fused 154. “Meckelian” groove 0= present 1= absent 155. Curvature of “Meckelian” groove (under tooth row) (applicable only to taxa with “Meckelian” groove) 0= parallel to 1= convergent on ventral border of mandible 156. “Coronoid” facet 0= present 1= absent 157. Vertical position of mandibular foramen 0= anteriorly placed, near back of dentition 1= near ventral margin, at root of angle 2= recessed dorsally from ventral margin, but below alveolar plane 3= recessed dorsally from ventral margin, at or above alveolar plane 158. Mandibular foramen dorsal to prominent longitudinal ridge 0= present 1= absent Skull – Rostrum 159. Septomaxilla 0= present 1= absent 160. Premaxilla, facial process dorsal extent 0= does not 1= does reach nasal 161. Premaxilla, facial process posterior extent 0= does not extend beyond canine 1= extends beyond canine but does not contact frontal 2= extends beyond canine and contacts frontal 162. Premaxilla, facial process with distinct finger‐like posterodorsal process 0= present 1= absent 163. Lateral margin of paracanine fossa 0= formed by maxilla =1 maxilla and premaxilla

164. Exit(s) of infraorbital canal 0= multiple 1= single 2= canal absent 165. Infraorbital foramen position 0= dorsal to ultimate premolar 1= to penultimate premolar or more anterior 2= to first molar or more posterior 166. Infraorbital canal length 0= long (more than one molar length) 1= short (less than one molar length (1). 167. Flaring of cheeks behind infraorbital foramen, seen in ventral view (Rougier et al., 1998: 83) 0= present 1= absent 168. Nasal 0= widest posteriorly 1= sides sub‐parallel 2= widest anteriorly 169. Nasal overhangs external nasal aperture 0= present 1= absent 170. Naso‐frontal suture with medial process of frontals wedged between nasals 0= present 1= absent 171. Naso‐frontal suture position 0= posterior to or even with 1= anterior to anterior orbital rim 172. Nasal foramina 0= present 1= absent 173. Frontal‐maxillary contact on rostrum 0= absent 1= present 174. Maxillary process of frontal (anterior projection of frontal) 0= weak or absent 1= elongate and thin 175. Preorbital length relative to postorbital 0= less than one third total length 1= more than one third 176. Lacrimal 0= present 1= absent 177. Facial process of lacrimal 0= large, triangular and pointed anteriorly 1= small, rectangular or crescentic (1)

178. Lacrimal tubercle 0= present 1= absent 179. Lacrimal foramen exposed on face 0= present 1= absent 180. Lacrimal foramen number 0= two 1= one 181. Lacrimal foramen within lacrimal 0= present 1= absent, with maxillary contribution 2= absent, with jugal contribution 182. Translacrimal canal 0= absent 1= present Skull – Palate 183. Premaxilla, palatal process 0= does not 1= does reach nearly or to canine alveolus 184. Premaxillary‐maxillary suture on palate 0= transverse 1= wedge‐shaped, pointing anteriorly 2= wedge‐shaped, pointing posteriorly 185. Incisive foramina 0= small, length of 1 or 2 incisors 1= intermediate, length of 3 or 4 incisors 2= elongate, more than half the palate length 186. Incisive foramina composition 0= between premaxilla and maxilla 1= within premaxilla 187. Palatal vacuities 0= absent 1= present 188. Major palatine foramen 0= within palatine 1= between palatine and maxilla 2= within maxilla 3= multiple small foramina 4= absent 189. Anterior extent of palatine on palate 0= to level of first molar 1= more posterior 2= more anterior 190. Palatal expansion with regard to ultimate molar

0= even with 1= posterior 2= anterior 191. Postpalatine torus 0= absent 1= present 192. Posterior nasal spine 0= weak or absent 1= prominent 193. Minor palatine foramen 0= small 1= large, with thin, posterior bony bridge 2= multiple small foramina 3‐ absent 194. Minor palatine foramen composition 0= palatine or maxilla‐palatine 1= palatine‐pterygoid 195. Maxilla with large shelf‐like expansion posterior to ultimate molar 0= absent 1= present Skull – Zygoma 196. Posterior edge of anterior zygomatic root 0= aligned with last molar 1= with anterior molars 2= with premolars 197. Zygomatic process of maxilla 0= present 1= vestigial 198. Jugal 0= present 1= absent 199. Jugal 0= contributes to anteroventral orbit and zygoma 1= contributes to zygoma 200. Maxillary‐jugal contact bifurcated 0= absent 1= present 201. Jugal‐lacrimal contact 0= present 1= absenT 202. Zygomatic arch 0= stout 1= delicate 2= incomplete

Skull – Orbit 203. Roots of molars exposed in orbit floor 0= absent 1= present 204. Palatine reaches infraorbital canal 0= present 1= absent 205. Lacrimal contributes to maxillary foramen 0= present 1= absent 206. Groove connects maxillary and sphenopalatine foramina 0= absent 1= present 207. Sphenopalatine foramen 0= within palatine 1= between palatine and maxilla 2= between palatine, maxilla, and frontal 3= within maxilla 208. Sphenopalatine foramen proximal to maxillary foramen 0= absent 1= present 209. Maxilla excluded from medial orbital wall 0= present 1= absent 210. Frontal and maxilla contact in medial orbital wall 0= absent 1= present 211. Orbital process of palatine 0= present 1= absent or with thin sliver in ventromedial wall of orbit 212. Ethmoid exposure in medial orbital wall 0= absent 1= present 213. Ethmoid foramen 0= between frontal and orbitosphenoid 1= within frontal 214. Foramen for frontal diploic vein 0= absent 1= present 215. Frontal foramen on skull roof 0= absent 1= present 216. Postorbital process 0= present, prominent 1= present, weak

2= absent 217. Postorbital process composition 0= frontal 1= parietal 218. Postorbital bar 0= absent 1= present 219. Dorsal process of jugal 0= weak or absent 1= strong 220. Optic foramen 0= absent 1= present 221. Optic foramen position 0= narrowly 1= broadly separated from sphenorbital fissure 2= not visible in lateral view 222. Orbitosphenoid 0= expanded anteriorly from optic foramen (or with anterior process for forms without optic foramen) 1= expanded dorsally from optic foramen (or with dorsal process for forms without optic foramen) 2= not expanded anteriorly or dorsally 223. Suboptic foramen 0= absent 1= present 224. Orbitotemporal canal 0= present 1= absent 225. Frontal/alisphenoid contact 0= dorsal plate of the alisphenoid contacting frontal at anterior corner 1= with more extensive contact with frontal (~50% of its dorsal border) 2= absent Skull – Braincase 226. Frontal length on midline 0= subequal to slightly smaller than parietal 1= less than half that of parietal 2= more than 50% longer than parietal 227. Frontoparietal suture 0= transverse 1= with anterior process of parietal off the midline 2= with anterior process of parietal on the midline 228. Temporal lines meet on midline to form sagittal crest 0= present 1= absent

229. Interparietal 0= absent 1= present 230. Nuchal crest 0= level with or anterior to foramen magnum 1= posterior to foramen magnum 231. Anterior lamina exposure on lateral braincase wall 0= present 1= absent 232. Squama of squamosal 0= absent 1= present 233. Foramina for temporal rami 0= on petrosal 1= on parietal and/or squama of squamosal 2= absent Skull – Mesocranium 234. Choanae 0= as wide as posterior palate 1= narrower 235. Vomer contacts pterygoid 0= absent 1= present 236. Pterygoids contact on midline 0= present 1= absent 237. Pterygopalatine crests 0= present 1= absent 238. Midline crest in basipharyngeal canal 0= absent 1= present 239. Entopterygoid process 0= absent 1= ends at anterior basisphenoid 2= approaches ear region 240. Midline rod‐shaped eminence on basisphenoid 0= absent 1= present 241. Ectopterygoid process of alisphenoid 0= absent 1= ends at anterior basisphenoid 2= approaches ear region 242. Ectopterygoid process of alisphenoid extent 0= long crest

1= narrow process (1) 243. Transverse canal foramen 0= absent 1= present 244. Exit for maxillary nerve relative to alisphenoid 0= behind 1= within 2= in front 245. Number of exit(s) for the mandibular branch of the trigeminal nerve 0= two 1= one 246. Foramen ovale composition 0= in petrosal (anterior lamina) 1= between petrosal and alisphenoid 2= in alisphenoid 3= between alisphenoid and squamosal 247. Foramen ovale position 0= on lateral wall of braincase 1= on ventral surface of skull 248. Alisphenoid canal 0= absent 1= present 249. Posterior opening of alisphenoid canal 0= separated from foramen ovale 1= in common depression with foramen ovale (1) Skull – Basicranium 250. Position of jaw articulation relative to fenestra vestibuli 0= at same level 1= in front 251. Glenoid fossa position 0= on zygoma 1= partially on braincase 252. Glenoid fossa shape 0= concave, open anteriorly 1= trough‐like 2= anteroposteriorly elongate 3= anteroposteriorly short 4= convex, open anteriorly 253. Glenoid fossa position relative to sphenoid on midline skull base 0= even with 1= higher 254. Glenoid process of jugal 0= present, with articular facet 1= present, without facet 2= absent

255. Glenoid process of alisphenoid 0= absent 1= present 256. Postglenoid process 0= absent 1= present 257. Postglenoid foramen 0= absent 1= present 258. Postglenoid foramen position 0= behind postglenoid process 1= medial or anterior to postglenoid process 2= on lateral aspect of braincase 259. Postglenoid foramen composition 0= within squamosal 1= behind squamosal 260. Suprameatal foramen 0= absent 1= present 261. Entoglenoid process of squamosal 0= absent 1= present, separate from postglenoid process 2= present, continuous with postglenoid process 262. Posttympanic crest of squamosal 0= absent 1= present 263. Carotid foramen 0= within basisphenoid 1= between basisphenoid and petrosal 2= absent 264. Cavum epiptericum 0= floored by petrosal 1= petrosal and alisphenoid 2= primarily or exclusively squamosal 3= primarily open as piriform fenestra 265. Alisphenoid tympanic process 0= absent 1= present 266. Basisphenoid tympanic process 0= absent 1= present 267. Basicochlear fissure 0= closed 1= patent

268. Epitympanic wing medial to promontorium 0= absent 1= flat 2= thickened 269. Rostral tympanic process of petrosal, on posteromedial aspect of promontorium 0= absent or low ridge 1= moderate ridge, contributing to posterodorsomedial bulla 2= tall ridge, contributing to ventral bulla 270. Course of internal carotid artery 0= lateral (transpromontorial) 1= medial (perbullar or extrabullar) 2= absent 271. Intratympanic vascular canal (for transpromontorial internal carotid) 0= absent 1= present 272. Deep groove for internal carotid artery excavated on anterior pole of promontorium 0= absent 1= present 273. Perbullar carotid canal (for medial internal carotid) 0= absent 1= present 274. Stapedial artery on promontorium 0= sulcus 1= canal 2= absent 275. Stapedial ratio 0= rounded, less than 1.8 1= elliptical, more than 1.8 276. Coiling of cochlea 0= less than 360° 1= 360° or greater 277. Pars cochlearis length 0= more than 13% of skull length 1= less than 10% of skull length 278. Promontorium shape 0= flat 1= globose 279. Promontorium depth relative to basioccipital 0= even with or ventral to 1= dorsal to 280. Intratympanic course of facial nerve 0= open in sulcus 1= open anteriorly, canal posteriorly 2= in canal

281. Tympanic aperture of hiatus Fallopii 0= in roof through petrosal 1= at anterior edge of petrosal 2= absent 3= via fenestra semilunaris 282. Prootic canal 0= present 1= absent 283. Prootic canal length and orientation 0= long and vertical 1= short and vertical 2= short and horizontal 284. Lateral flange 0= parallels length of promontorium 1= greatly reduced or absent 285. Length of bony shelf lateral to promonotorium (lateral trough or tegmen tympani) 0= extended anteriorly as far as promontorium 1= confined posterolaterally 2= prolonged anterior to promontorium 286. Width of bony shelf lateral to promonotorium (lateral trough or tegmen tympani) 0= uniform 1= expanded anteriorly 287. Inflation of bony shelf lateral to promontorium (lateral trough or tegmen tympani) 0= absent 1= present 288. Stapedial canal on bony shelf lateral to promontorium (lateral trough or tegmen tympani) 0= absent 1= present 289. Tensor tympani fossa 0= shallow 1= deep circular pit 290. Medial process of squamosal in tympanic cavity 0= absent 1= present 291. Hypotympanic sinus 0= absent 1= formed by squamosal, petrosal, and alisphenoid 2= formed by alisphenoid and petrosal 3= formed by petrosal 292. Epitympanic recess/fossa incudis size 0= subequal 1= epitympanic recess larger 2= no visible depression for epitympanic recess

293. Epitympanic recess lateral wall 0= with small contribution to posterolateral wall by squamosal 1= with extensive contribution to lateral wall by squamosal 2= with no squamosal contribution 294. Fossa incudis (Rougier et al., 1998: 137) 0= continuous with 1= separated from epitympanic recess 295. Floor ventral to fossa incudis 0= absent 1= formed by squamosal 2= formed by ectotympanic 296. Fossa incudis position relative to fenestra vestibuli 0= lateral 1= anterior 297. Foramen for ramus superior of stapedial artery 0= on petrosal 1= on petrosal‐squamosal suture 2= absent 298. Position of ramus superior foramen relative to fenestra vestibuli 0= posterior or lateral 1= anterior 299. Ascending canal 0= intramural 1= intracranial 2= absent 300. Stapedius fossa 0= twice the size of fenestra vestibuli 1= small and shallow 301. Cochlear canaliculus visible canal in middle ear space 0= absent 1= present 302. Cochlear fossula 0= weak or absent 1= distinct pit behind fenestra cochleae 303. Fenestra cochleae position to fenestra vestibuli 0= posteromedial 1= posterior 304. Posterior septum shields fenestra cochleae 0= absent 1= present 305. Paroccipital process (sensu Wible and Hopson, 1993) 0= vertical 1= slanted, projecting anteroventrally as flange towards back of promontorium 2= indistinct to absent

306. Caudal tympanic process of petrosal notched 0= absent 1= present 307. Crista interfenestralis and caudal tympanic process of the petrosal connected by curved ridge 0= absent 1= present 308. “Tympanic process” 0= absent 1= present, low 2= present, high. 309. “Tympanic process” composition 0= petrosal 1= petrosal and exoccipital 310. Rear margin of auditory region 0= marked by steep wall 1= extended onto a flat surface 311. Inferior petrosal sinus 0= intrapetrosal 1= between petrosal, basisphenoid, and basioccipital 2= endocranial 312. Jugular foramen size relative to fenestra cochleae 0= subequal 1= larger 313. Jugular foramen 0= confluent with 1= separated from opening for inferior petrosal sinus 314. Hypoglossal foramen 0= two or more 1= one 315. Hypoglossal foramen housed in opening larger than jugular foramen 0= absent 1= present 316. Paracondylar (“paroccipital”) process of exoccipital (sensu Evans and Christensen, 1979) 0= weak or absent 1= prominent, vertical 2= prominent, posteriorly directed 317. Ectotympanic 0= phaneric or visible in ventral view 1= aphaneric or hidden by auditory bulla 318. Ectotympanic shape 0= ring‐like 1= fusiform 2= expanded

319. Anterior crus of ectotympanic broadly contacts facet on squamosal 0= absent 1= present 320. Elongate ossified external acoustic canal 0= absent 1= present 321. Roof of external acoustic meatus 0= petrosal 1= squamosal 322. Entotympanic 0= absent 1= present 323. Pit on ectotympanic for hyoid 0= absent 1= present 324. Hyoid arch contributes to bulla 0= absent 1= present 325. Dorsum sellae 0= tall 1= low 326. Posterior clinoid process contacts anterior pole of promontorium 0= absent 1= present 327. Position of sulcus for anterior distributary of transverse sinus relative to subarcuate fossa 0= anterolateral 1= posterolateral 328. Wall separating cavum supracochleare from cavum epiptericum 0= absent 1= incomplete, with fenestra semilunaris 2= complete 329. Crista petrosa 0= vestigial or absent 1= tall, thin crest 330. Subarcuate fossa aperture 0= not constricted 1= constricted 2= fossa absent 331. Anterior semicircular canal 0= does 1= does not form lateral wall of subarcuate fossa aperture 332. Internal acoustic meatus 0= deep, with thick prefacial commissure 1= shallow, with thin prefacial commissure

Skull – Occiput 333. Posttemporal canal 0= large 1= small 2= absent 334. Posttemporal canal composition 0= between petrosal and squamosal 1= within petrosal 335. Posttemporal canal position 0= on occiput 1= dorsal to external acoustic meatus 336. Mastoid foramen 0= absent 1= two in mastoid 2= one in mastoid 3= between mastoid and supraoccipital 337. Amastoidy or lack of occipital exposure of mastoid 0= absent 1= present 338. Dorsal margin of foramen magnum 0= formed by exoccipitals 1= by exoccipitals and supraoccipital

Matrix for ancient placement of Taeniodonta Montanalestes 00[01]????[012]????????????????????[01]?1????????????????????1001100011 10??????????????????????????????????????????01000100111021111000000[12] 3010021201100201010110[012]01- 021???????????????????????????????????????????????????????????????????? ??????????????????????????????????????????????????????????????????????? ????????????????????????????????????????? Eozhelestes 000????0??????????0?????11??0?????????????????000[01]?1????????1?0????? ?????????????????????????????????????000001001110223020001?0?2????????? ???????????0[01]0?????????????????????????????????????????????????????? ??????????????????????????????????????????????????????????????????????? ???????????????????????????????????????????????????????????? Cimolestes 001[03]00?1??????021?001001010?1?1?10-10012[01][12]0000010- - 000000000100110010210022001011001111221120101[01][01]000010000000000111 0211120[01]00001[12]01001120010????000?00[01]01- 121????11[01]?10??0????01?110???????2010??01???????????????????10- ??????????????????????????????????????????????????????????????????????? ????????????????????????????????????????????????? Kennalestes 001000[012]1?00001000?0?00000000101000-10011121100001-- 00000101010021202120002200101100111112112001112000010000000101111021111 0000001201002120010020000??10001-1211100011010?0?10- 101111000???0000101100?0000110?10000000??2--

?10100???0001111?01?2?20?11300-1000?01110121?3000101-000111100?1- 10000000?1?01??[01]00100211101100110?1?00000?00211112--20? Asioryctes 00100000000001000?000000000?1?1000-20011220000001-- 1001010101002100102100220010110011111[02]112001100-- 001000000020111102111100000001010021200100200010010001- 1211100011010?0010-1001110000000??010110000010110?10000000??2-- 010100[02]100001111?0111020011300-1000101110121?3000101-00011110001- 10000000?1101??[01]0010021110110011001?00000?0??????2--101 Zalambdalestes 00[12][01]00[23]11110?21211211000111?[12]?1000- 21111120001000--00012100010022002021002121111102101100-22001200-- 00110001002211131223121000001101011120[01]100200001010001- 121110001100000110- 1000100101000200[01]011010000011101200110100000010100101[01]00111100111 120011300-1000[12]0111012103000101-00011110011- 100000000101111[01]00100211111[12]0000001101000?102???1100101 Barunlestes 002[01]00?1??????1211211011111?2?1----201111200?000--- 0??12100010022002021002121111102101100-22001200-- 0011000100221113122312100001-1010111200100200001010101- 121???0?11000????0- 100?110?0???0220101001?00????1?1[13]01100??0????10?0??????011?1??111120 01130??1000?0111012103000101-0?011110011- 10000000?1?11???00100211111?00000????1?0??1??????????01 Alveugena 0010?13??????0????????11010?1?201?-100110100?0010-- 000000?0?100120020010020001001001??322??2101?210000110000?-00- 31?0111?20000?0120????????????????????????????????????????????????????? ??????????????????????????????????????????????????????????????????????? ??????????????????????????????????????????????????????????????????????? ???????????? Schowalteria 001?0??2?1???01?00??0011010?1?2010-??00??2???0011-- 000001?011001200200100- 00??????????3??????0??1100?????100???1??1?001?????0?10220100???10?????? ??????2?1- 1???????11??????????????????????????????1?00002???????????????????????? ??????????????????????????????????????????????????????????????????????? ?????????????????????????????????????????

Matrix for modern placement of Taeniodonta Montanalestes 00[01]????[012]????????????????????[01]?1????????????????????1001100011 10??????????????????????????????????????????01000100111021111000000[12] 3010021201100201010110[012]01- 021???????????????????????????????????????????????????????????????????? ??????????????????????????????????????????????????????????????????????? ????????????????????????????????????????? Cimolestes 001[03]00?1??????021?001001010?1?1?10-10012[01][12]0000010- - 000000000100110010210022001011001111221120101[01][01]000010000000000111 0211120[01]00001[12]01001120010????000?00[01]01- 121????11[01]?10??0????01?110???????2010??01???????????????????10- ??????????????????????????????????????????????????????????????????????? ?????????????????????????????????????????????????

100

Vulpavus 00[12]000210000000000000001010?[12]?0111-00011230000011-- 00000000?12022002021002222111102111100- 22101111[01]10011001100210111122131000010[12][12]0?0011?00?0000001112?? 01- 1??11[01]011001[12]010110001111001?0?01[02]?????01?0010[01]?0?????????? 0000110?01?01001111??110?0???2120111000?01100?011300000000-0?11010?1- 12000102[01]1[01]0010[01]000000100-1[12]10102???01?00????????????0? Miacis 001000210000000000000?01010?1?0111-0000123000101?-- 00010000012022002021002221011102111100- 221001[02][01]01001100110010011111023100001[01]?10100?10000?0????1111?? ?1- 1[123]1?????10012?10?10???????????????20???0???????10?1000000110000?10? 010010?111111?101011?21[23]0111000?011000111[13]00000000-0111010?1- 1200011111000???00000010101[12]10002???01?00????????????0? Diacodexis 00100021?0000?0400000001110?1?2001-20012001100011-- 000201001111120020210022221111021111221221022230000100000?0210111222212 0001002[12]010011000000300001[01]2000?????1110111010?0011010101100????? ??1103-03000000??002001001110000100000000011111??1?1010021200- 1000101100???1300000000-???1?1??1- ??0?01??????????????02????1?11[01]0102?01000????????2--30? Ptilocercus 0020003110000021100010101100200000-10011020100----- 00022100110012202021002212211100110100- 2211112201001000001011021312231200010031010011200000200001020001- 13111200101120001110011010010000000113001000111111100100011000111010020 1110111011101010021201010000011001101100000010-111110211- 1210100211100[01]11100002000-120110010101100100211112--301 Tribosphenomys 0032114301221-121131-02-2--?3-2----0002???????------??0210-010122002021002222211122-101220221022031-001111----- 21313122102010001-2100?102100?02??0???10101-131?1??- 1110???????????????- ????????????10?????1??????????????????????????????????????????????????? ??????????????????????????????????????????????????????????????????????? ????????????? Rhombomylus 0032114301221-121131-02-2--?3-2----2[01]0?2[01]10200----- 011220200111233-----0--021201-22-20100-221022032-1?1111----- 110131211120000003[12]100011210000210000020101-1211120- 1110200011000111110-0100002003- 01000010111010110010000011010121200011111111202002120[01]01021200110000 2[13]000102---211110[12]11-12100103120202-2?01002000- 1210[01]0202111000000201012--201 Potamogale 0023002101200023000010101100202000-20012210000----- 11012001110021102022102212001100111100-22000100-- 0010000000211111020032-0000013010020000000000001010201- 131111011210111011111--111-1000032100000001---210-00100001002--- 1001101101011110110100-02120111100-0110?0211211101000-011100011- 1100001111010111001000102012101000110100110020111100001 Rhynchocyon 00210051-----103100011001000102001-12011220201001-- 11[01]221200222033-----0--021211-02-10100-22101100--0-1-- 000011103122220-2-000-012011100000000300101020201- 13111001100110111101001110012000021010010000101000001000001001001011102 0000112101101011021200-104020010101011110?1010-111100031- 101010011202?011?0???10?201[12]?100001111101100211112--201 101

Alveugena 0010?13??????0????????11010?1?201?-100110100?0010-- 000000?0?100120020010020001001001??322??2101?210000110000?-00- 31?0111?20000?0120????????????????????????????????????????????????????? ??????????????????????????????????????????????????????????????????????? ??????????????????????????????????????????????????????????????????????? ???????????? Schowalteria 001?0??2?1???01?00??0011010?1?2010-??00??2???0011-- 000001?011001200200100- 00??????????3??????0??1100?????100???1??1?001?????0?10220100???10?????? ??????2?1- 1???????11??????????????????????????????1?00002???????????????????????? ??????????????????????????????????????????????????????????????????????? ?????????????????????????????????????????

102