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An Investigation of the Phylogenetic Affinities of Sivaladapidae within Adapoidea

Kathleen Rust CUNY Hunter College

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This work is made publicly available by the City University of New York (CUNY). Contact: [email protected] An Investigation of the Phylogenetic Affinities of Sivaladapidae within Adapoidea

by

Kathleen L. Rust

Submitted in partial fulfillment of the requirements for the degree of Master of Arts Anthropology, Hunter College The City University of New York

2018

Thesis Sponsor:

April 23, 2018 Dr. Chris Gilbert Date Signature

April 23, 2018 Dr. Stephen Chester Date Signature of Second Reader

Table of Contents Abstract ...... 3 Introduction ...... 4 Background ...... 6 Adapoid and Sivaladapid Morphology ...... 6 History, Discovery, and Debate ...... 11 Material and Methods ...... 20 Results ...... 24 Discussion ...... 27 Conclusion ...... 32 Acknowledgments...... 33 Tables and Figures ...... 34 Figures Figure 1 ...... 34 Figure 2 ...... 35 Figure 3 ...... 36 Figure 4 ...... 37 Figure 5 ...... 39 Figure 6 ...... 40 Figure 7 ...... 41 Figure 8 ...... 42 Figure 9 ...... 48 Figure 10 ...... 49 Figure 11 ...... 50 Figure 12 ...... 51 Tables Table 1 ...... 38 Table 2 ...... 43 Table 3 ...... 52 Table 4 ...... 61 Table 5 ...... 68 Table 6 ...... 72 Table 7 ...... 75 Table 8 ...... 79 Table 9 ...... 81 Table 10 ...... 85 Table 11 ...... 87 Table 12 ...... 91 Table 13 ...... 92 Table 14 ...... 93 References ...... 94

2 Abstract

Research on the earliest fossil primates has deepened our understanding of primate evolutionary history while simultaneously generating competing hypotheses regarding the phylogenetic affinities of adapoids and omomyoids. This study presents a phylogenetic analysis incorporating 82 dental characters that aims to clarify the evolutionary relationships of the sivaladapids within the broader context of Adapoidea. Previous hypotheses have suggested that sivaladapids are closely related to a European adapoid clade (Qi and Beard, 1998), or that they share a close evolutionary relationship with an Asian adapoid clade recognized as the asiadapids

(Rose et al., 2009; Godinot, 2015). Results here suggest that sivaladapids share a closer evolutionary relationship with European adapoids rather than asiadapids, but that the relationships among the major families of Eurasian adapoids are probably closely intertwined with numerous biogeographic connections between the two continents during the Eocene.

Future investigations examining additional specimens and postcranial characters will be necessary to offer further resolution of the specific relationships of Sivaladapidae and their origins.

3 Introduction

For almost 150 years, the fossil record has informed studies of the earliest primates

(Cope, 1872a; 1872b; Gregory, 1920; Gregory et al., 1938; Simpson, 1940; Szalay and Delson

1979, Fleagle, 2013). Fossils of the first primates shed light on the evolutionary history of the

Primate Order, fueling several competing hypotheses of their phylogenetic relationships to extant strepsirrhines and haplorhines (Gregory, 1920; Hofstetter, 1977; Gingerich and Simons, 1977;

Gingerich, 1977a, 1977b; Cartmill and Kay, 1978; Rasmussen, 1986; Seiffert at el., 2009;

Gingerich, 2012; Ni et al., 2013). In recent years, invigorated paleontological efforts across the globe have identified new species and contributed to a growing fossil record that illustrates the morphological diversity and phylogenetic complexity of the earliest primates (Marivaux et al.,

2002; Ni et al., 2004, 2016; Beard et al., 2007; Franzen et al., 2009; Rose et al., 2009; Gebo et al., 2012; Dunn et al., 2016; Boyer et al., 2017; Femenias-Gaul et al., 2017; Gilbert et al., 2017;

Seiffert et al., 2018). The study presented here contributes to the understanding of the evolutionary relationships among the earliest known primates through a phylogenetic analysis that focuses on the group of adapoids placed in the family Sivaladapidae, or sivaladapids.

Primates of modern aspect (Hoffstetter, 1977) first appear in the fossil record approximately in the early Eocene, following the Cretaceous-Paleogene mass extinction event that includes the demise of non-avian dinosaurs (Rose, 1995; Godinot, 1998; Fleagle, 2013;

Godinot, 2015). Unlike the plesiadapiforms, these early primates all possess the morphological synapomorphies that characterize extant primates, including a petrosal auditory bulla, a postorbital bar, orbital convergence, a maximum of two incisors, an opposable hallux, increased brain size, and nails instead of claws on most digits (Le Gros Clark, 1959; Martin, 1968; Rose,

1995; Rose, 2006; Fleagle, 2013; Godinot, 2015; Silcox et al., 2015; Gilbert and Jungers, 2017).

4 It was during the early Eocene that several mammalian orders and the earliest primates, broadly recognized at the infraorder level as Adapiformes and Omomyiformes, began to diversify.

Adapiforms and omomyiforms (hereafter discussed at the superfamily level as adapoids and omomyoids) have traditionally been described as lemur-like and tarsier-like, respectively (Rose,

1995). Primates from these two groups appear simultaneously in Asia, Europe, and North

America approximately 56 Ma (e.g., Figure 5) (Rose, 1995; Fleagle, 2013; Godinot, 2015).

Their migration between and across these northern continents was facilitated by the Paleocene-

Eocene Thermal Maximum, a dramatic and rapid global warming event that occurred at the

Paleocene-Eocene boundary (Smith et al., 2006; Beard, 2008). The warmer climatic conditions enabled the successful radiation and diversification of adapoids and omomyoids for several million years.

By the end of the Eocene and beginning of the Oligocene, most species of adapoids disappear from the fossil record in North America and Europe as the climate changed (Gingerich and Sahni, 1984; Qi and Beard, 1998; Fleagle, 2013). One distinct group of adapoids, recognized today as sivaladapids, survived the changing temperatures and persisted late into the

Miocene of Asia. Their ability to weather the changing climates has been related to their biogeographic distribution across Asia, affording them easy migration into the tropical refugia of southern and southeastern Asia (Qi and Beard, 1998). Gingerich and Sahni (1984) suggest their extinction by the Late Miocene may have been associated with the immigration of colobine monkeys, known arboreal folivores that may have outcompeted the sivaladapids for resources.

Today approximately 46 genera and 105 species of fossil adapoids are recognized

(Godinot, 2015). Sivaladapids, compared to other adapoid taxa, are still poorly known in the fossil record, with no good cranial material yet described. They are instead represented by

5 partial dentaries, isolated teeth, and limited postcrania. Understanding of the phylogenetic affinities of Sivaladapidae and the broader evolutionary relationships within Adapoidea has been met with competing hypotheses (Qi and Beard, 1998; Rose et al., 2009; Godinot, 2015); unequal representation of anatomical elements typical of fossil taxa further hinders systematic resolution within and among adapoid families. The continued disagreement of adapoid systematics has been accompanied by different taxonomic compositions of the superfamily. Here, for convenience, adapoids are recognized taxonomically following Fleagle (2013), who identifies six distinct families: Notharctidae, Cercamoniidae, Asiadapidae, Caenopithecidae, Adapidae, and

Sivaladapidae.

Given the complicated and debated nature of adapoid systematics, this study seeks to investigate the phylogenetic affinities of sivaladapids relative to other adapoid families. While previous research on sivaladapids has focused on relationships within the family (Qi and Beard,

1998; Gilbert et al., 2017), or have aimed to more broadly clarify adapoid relationships within the larger primate radiation (e.g., Ni et al., 2016), this study analyzes sivaladapids within the broader context of Adapoidea. The phylogenetic analysis here attempts to clarify relationships of sivaladapids within Adapoidea to understand the origins of Sivaladapidae.

Background

Adapoid and Sivaladapid Morphology

Adapoids, along with omomyoids can generally be characterized as the most primitive primates (Szalay and Delson, 1979; Rose, 1995; Fleagle, 2013; Godinot, 2015). Adapoids are known by many thousands of isolated teeth, hundreds of dentaries, numerous skulls, and a few nearly complete skeletons (Rose, 2006). They are best represented in the Eocene fossil record of

North America, spanning from northern localities of Wyoming to as far south as Texas. Their

6 anatomy bears resemblance in many aspects to extant strepsirrhines, and they have been traditionally referred to as “lemur-like” in the literature (Rose, 1995, 2006; Fleagle, 2013;

Godinot, 2015). Some researchers have critiqued this comparison, arguing that the adaptive diversity of extant strepsirrhines is too broad to adequately characterize adapoids; while certain anatomical characteristics of adapoids do bear similarity to strepsirrhines, the morphological and adaptive diversity of extant strepsirrhines offers little specificity in understanding the biological profile of extinct adapoids (Covert, 2002; Covert and Williams, 1994). However, with discovery of additional adapoid fossils and newly identified genera and species, it could be argued that the extinct clade does match the adaptive diversity that exists in extant strepsirrhines, in that the diversity of fossil adapoids fits well within the confines of the comparison to strepsirrhines.

Concerning their phylogenetic affinities to extant primates, it has been suggested that the adapoids gave rise to modern strepsirrhines and haplorhines (Gingerich, 1975; Gingerich, 1977a,

1977b; Gingerich, 2012), are stem anthropoids (Rasmussen, 1994), are stem strepsirrhines

(Beard et al., 1988; Seiffert, 2005), or may have a sister group relationship with crown strepsirrhines (Rosenberger et al., 1985; Seiffert, 2009). These conflicting hypotheses have generated debate, warranting close consideration of the synapomorphies that have been proposed to support these hypotheses, respectively. Those that do not view adapoids as stem strepsirrhines argue that the synapomorphies characterizing the living lemuriforms are all soft-tissue features and unidentifiable in the fossil record, including a wet nose with lateral slits, a frenulum, and philtrum (Rasmussun & Nekaris, 1998); however, numerous authors demonstrate that fossil adapoids bear hard-part morphological features that corroborate the presence of a wet rhinarium in the extinct species, most notably in presence of a median-gap and the spacing, occlusion, and size-shape relationship of the upper incisors of notharctids, combined with a long and

7 horizontally-oriented nasolacrimal canal (Rosenberger et al., 1985; Rossie et al., 2006; Rossie and Smith, 2007; Tabuce et al., 2009; Kirk et al., 2014). The authors argue that these hard-part features directly correlate with the soft-tissue synapomorphies that typify the extant lemuriforms, and thus comparison of strepsirrhines and adapoids is duly justified. While these characters are shared among adapoids and extant strepsirrhines, others have pointed out that they are symplesiomorphies of the primate order and shed no light on the particular phylogenetic affinities of adapoids (Rasmussen and Nekaris, 1998). Conversely, Covert and Williams (1994) and Beard et al. (1988) have also argued that other hard-part morphological features found in the postcrania of adapoid fossils are indeed true synapomorphies shared between adapoids and strepsirrhines; these include a sloping talofibular-joint facet, a laterally positioned groove for the flexor hallucis longus relative to the posterior portion of the tibiotalar joint, a large posterior trochlear shelf, contiguous navicular cuboid articulation with the mesocuneiform and the ectocuneiform, and an anteriolateraly rotated medial malleolus of the tibia (Dagosto, 1993, 2007;

Dagosto and Gebo, 1994). Boyer et al. (2013) also highlight similarities in metacarpal morphology shared between the Eocene adapoids and extant strepsirrhines. Significantly, many if not all adapoids appear to have a grooming claw or nail present on their 2nd pedal digit when preserved, another likely synapomorphy shared with extant strepsirrhines (Maiolino et al., 2012; von Koenigswald et al., 2012; Gilbert and Maiolino, 2015).

While the earliest adapoids share common dental morphology with the contemporaneous omomyoids, they are distinguished by a number of characters in both the dentition, and in particular, the broader skull and postcranium. The primitive dental formula of adapoids (and omomyoids) is 2.1.4.3/2.1.4.3, a condition that is retained by several adapoid lineages. They possess small and most often vertically implanted incisors that appear broad and spatulate-like,

8 with I2 typically larger than I1 (Rose, 1995, 2006; Gebo, 2002; Fleagle, 2013). Some species exhibit canine dimorphism, but it has been established that the manner in which the dimorphism is achieved in adapoids differs developmentally than that observed in anthropoids (Gingerich,

1995; Schwartz et al., 2005). The anterior premolars are simple and often caniniform while the posterior premolars can be semimolariform (Rose, 2006). The upper molars are typically broad, and certain lineages independently evolved a hypocone. The lower molars are long and narrow in shape but vary in cusp number and morphology among the adapoid families; some taxa exhibit low, rounded cusps, whereas others possess sharp crests and reduced or absent paraconids. Trigonids are typically small. The convergent evolution of shearing crests and mandibular fusion is documented in several lineages, hypothesized as adaptations to folivory and/or also related to scaling and increasing body size (Rose, 1995; Ravosa, 1996; Qi and Beard,

1998; Gebo, 2002; Rose, 2006; Fleagle, 2013). The cranial and postcranial morphology of adapoids are not as well represented in the fossil record as the dental morphology. The known skulls typically have long snouts, small orbits, and a large braincase compared to plesiadapiforms but smaller when compared to extant lemurs (Rose, 2006; Fleagle, 2013; Gilbert and Jungers, 2017). Adapoid skulls also display a suspended ectotympanic ring within the inflated auditory bulla, a condition that is also present in extant lemurs (Rose, 2006; Fleagle,

2013). Limb bones are similar to that of strepsirrhines but differ slightly in their robusticity

(Fleagle, 2013); certain fossils (e.g., adapids) display morphology indicating a slow, cautious arboreality like that of living lorises (Gebo et al., 2012), while others display limb proportions and morphology indicative of more active arboreality and some leaping (Dagosto, 1993, 2007;

Rose, 2006). Postcranial elements vary also in size, and body mass estimates from dental

9 regression analyses show a range of adapoid body size from 100g to 7kg, a range comparable to living lemurs (Covert, 2002; Fleagle, 2013).

Within Adapoidea, diagnostic features of sivaladapids derive mostly from dentition, as they are not as well represented anatomically in the fossil record compared to North American and European adapoids. Furthermore, the Miocene taxa are much better represented than the basal Eocene taxa. Dentally, sivaladapids are distinctive in their combination of strong lower molar hypoconulids, twinning of the lower hypoconulid with the entoconid accompanied by a distinct notch between the two cusps, and a strong, continuous lingual cingulum on the upper molars (Figures: 1-4) (Gebo, 2002; Godinot, 2015). The Eocene forms are distinct with the presence of hypocones and pericones that arise from the cingulum on the upper molars, whereas the Miocene forms appear to have secondarily lost these cusps (Figures: 1-2) (Qi and Beard,

1998). As a whole, the group is united by the loss of the first premolar, with a dental formula of

2.1.3.3, but there is variation in root number and shape of lower P2; a two-rooted P2 is observed in Hoanghonius and Wailkeia, differing from the single-rooted P2 in the Miocene species (Gebo,

2002). Sivaladapis, Sinoadapis, and Indraloris are similar in sharing highly-crested lower molars and prominent mesostyles and parastyles that accompany well-developed shearing crests on the upper molars, likely associated with a folivorous diet (Figure 4); this morphology is also expressed but to a lesser degree in Guangxilemur (Figure 3) (Gingerich and Sahni, 1979;

Gingerich and Sahni, 1984; Qi and Beard, 1998; Gebo, 2002). The earlier and more primitive hoanghoniines exhibit more bunodont, lower-crowned lower molars with a more distinct paraconid than the Miocene forms; the upper molars bear less developed shearing crests and lack stylar cusps (Gingerich et al., 1994). Our knowledge of sivaladapid postcranial morphology is limited, based on only a few elements representing Hoanghonius and some isolated postcrania

10 potentially assignable to taxa from the late Eocene Pondaung Formation (Beard et al., 2007;

Gebo et al., 2015). Additional sivaladapid postcranial elements described to date include a proximal metatarsal and a second pedal distal phalanx with morphology suggesting the presence of a grooming claw (Gebo et al., 1999; Gebo et al., 2015).

History, Discovery, and Debate

While sivaldapids were not recognized as distinct adapoid primates until the late 1970’s

(Gingerich and Sahni, 1979; Thomas and Verma, 1979), the first adapoid fossil, a severely crushed palate and right mandible from the late Eocene of France, was described by Cuvier in

1821. He identified the fossil as having artiodactyl affinities and named it Adapis parisiensis

(“ad-apis” – toward the sacred bull Apis) (Gingerich, 1975; Gebo, 2002). Caenopithecus lemuroides reserves the title as the first actual adapoid to be recognized as a primate; in 1862,

Rütimeyer referred to the dental remains from the locality of Egerkingen in Switzerland as a middle Eocene primate, a hard sell to his contemporaries (Gebo, 2002; Godinot, 2015). Years following, more complete primate fossil remains were discovered in North American and

European Eocene deposits; these discoveries and their morphological affinities to extant lemuriforms compelled the scientific community to accept a much older and more widely spread primate order than was previously hypothesized. Given the unequivocal fossil evidence of

Eocene primates, Adapis was recognized as a primate and by the twentieth century, was aligned with the Order Primates (Gebo, 2002; Godinot, 2015)

The history, discovery, and identification of the sivaladapids are also complex. Several of the Miocene species belonging to the family today had been previously misidentified or debated among researchers, affecting taxonomic identification and systematic placement of species that spanned several years in the literature (Tattersall, 1968; Gingerich and Sahni, 1979;

11 Szalay and Delson, 1979; Chopra and Vasishat, 1980; Gingerich and Sahni, 1984). It wasn’t until Gingerich and Sahni (1979) and then Thomas and Verma (1979) proposed Sivaladapinae as a new subfamily that Asian taxa were recognized as a distinct clade. Sivalapadis and Indraloris were the first species to comprise the subfamily. Fossils of Sivaladapis from Siwalik deposits of

India were originally described by Pilgrim (1932); he interpreted the partial mandible, with only one lower molar crown present, as possessing a maximum of two lower molars and subsequently assigned it to the carnivoran genus and species Sivanasua palaeindica (Gingerich and Sahni,

1979; Chopra and Vasishat, 1980; Gingerich and Sahni, 1984). During the Yale North India

Expedition of 1932 and 1933, Lewis recovered a single tooth from the Nagri horizon of the

Siwalik deposits surrounding Haritalyangar which he identified as a lorisid primate, assigning it a new genus and species, Indraloris lulli. He suggested it could be derived from Adapidae but lacked sufficient material to support the phylogenetic placement (Lewis, 1933). Years later,

Tattersall (1968) identified fossil material in the Yale Peabody Museum collections that he attributed to Indraloris, pointing out similarity between Indraloris and the Sivanasua material.

Based on his observations, he considered the affinity of Sivanasua to procyonid carnivorans to be incorrect (Tatersall, 1968; Chopra and Vasishat, 1980). The identification of additional

Sivanasua fossil specimens as possessing three (rather than two) lower molars allowed for the direct comparison to I. lulli. The comparison showed a close phylogenetic relationship based on shared morphology including hypoconulid-entoconid twinning, supporting the renaming

Sivanasua to Sivaladapis (Gingerich and Shani, 1979, 1984). With discovery of more complete specimens lacking morphological synapomorphies with extant lorisids (i.e., a toothcomb),

Indraloris and Sivaladapis were considered closely related adapoids and designated to the

12 subfamily Sivaladapinae, which was then later promoted to the Sivaladapidae (Thomas and

Verma, 1979; Gingerich and Sahni, 1979, 1984; Godinot, 1998).

While the discovery of additional Miocene adapoid fossils that displayed prominent upper molar shearing crests and hypoconulids twinned with entoconids on the lower molars provided overwhelming support for a distinct Asian clade, gaps in the fossil record left researchers wondering how the Middle and Late Miocene sivaladapids were connected to the larger adapoid radiation of the Eocene (Szalay and Delson, 1979; Qi and Beard, 1998). Some researchers had noted the morphological similarities between Eocene and Miocene taxa, but their potential phylogenetic connection had been either dismissed or not investigated given the lack of fossil evidence bridging the two clades (Ducrocq and Chaimanee 1995; Gingerich et al., 1994,

Qi and Beard, 1998). While adapoid fossils from China and Thailand were known from the

Eocene, the temporal and morphological connections to the Miocene forms were still missing.

For example, a Middle Eocene dentary containing M1 and M2 from Shanxi Province of China was described by Zdanksy (1930), who named the fossil Hoanghonius stehlini but classified it as incertae sedis. He noted that the morphology of the lower molars was similar to that of

Smilodectes, a confirmed Eocene primate of North America (Gingerich et al., 1994; Zdansky,

1930), but lacked sufficient material to confirm the systematic position of Hoanghonius.

Authors debated the placement Hoanghonius until Szalay (1974) and Gingerich (1975) recognized it as an adapoid. Following Zdanksy’s (1930) original description, additional Late

Eocene fossil specimens were discovered from nearby localities and assigned to the genus (Woo and Chow, 1957). Gingerich et al. (1994) examined the specimens recovered by Woo and Chow

(1957) and observed morphological differences between the more recently recovered fossil and the original Zdanksy (1930) material, thus warranting a new genus and species for the dental

13 specimens described as Hoanghonius by Woo and Chow (1957). Gingerich et al. (1994) renamed the referred material as Rencunius zhoui, assigning it and Hoanghonius to a new subfamily, Hoanghoniinae. Support for a new subfamily came from a combination of shared morphology between Rencunius and Hoanghonius not seen in other adapoid taxa, including strong pericones and strong hypocones on the lingual cingulum of the upper molars, along with hypoconulid-entoconid twinning (Figures 1-2) (Gingerich et al., 1994).

Recovery of Eocene primate fossils of similar morphology in Thailand offered further support for Hoanghoniinae as a distinct Asian Eocene adapoid subfamily. Ducrocq and

Chaimanee (1995) described a lower dentary from the Eocene Krabi Basin of Southern Thailand as an anthropoid, naming it Wailekia orientale. Authors later recognized the overall similarity of

Wailekia to Hoanghonius, exhibiting lower molars with a twinned entoconid and hypoconulid and lacking a paraconid. Based on their shared morphology, Qi and Beard (1998), Godinot

(1998), and Gingerich et al. (1994) advocated for the placement of the Thailand adapoid in

Hoanghoniinae.

Finally, recovery an upper M2 displaying intermediate morphology from the Late Eocene

Gongkang Formation in southern China offered some phylogenetic resolution between the

Miocene sivaladapids and Eocene Hoanghoniinae (Qi and Beard, 1998). This temporally and morphologically intermediate upper molar, recognized as Guangxilemur tongi, bridged the gap between the Miocene and Eocene forms with its unique combination of dental characters, possessing certain features present in Hoanghonius and more derived traits present in Miocene sivaladapids. For example, the M2 of Guangxilemur possesses a pericone and hypocone, characters shared with Hoanghonius and Rencunius while appearing simultaneously advanced in its well-developed and slightly W-shaped external shearing crest, a derived trait shared with the

14 Miocene sivaladapids (Figure 3). Phylogenetic analysis by Qi and Beard (1998) supported the close relationship of the Eocene hoanghoniines and the Miocene sivaladapines via

Guangxilemur, placing G. tongi directly between the two subfamilies. Further corroborating the transitional stage of Guangxilemur, Marivaux et al. (2002) described isolated lower molars from

Oligocene deposits in Pakistan displaying intermediate morphology; the maxillary and mandibular dental specimens were assigned to the genus and identified as a new species, named

Guangxilemur singsilai.

The additional Guangxilemur material described by Marivaux et al. (2002) not only reinforced the intermediate phylogenetic position of the genus, but it also allowed for direct comparison of the lower dentition between the Miocene and Eocene taxa. The lower molars exhibit a quasi-indistinct paraconid and the lingually positioned hypoconulid twinned with the entoconid. The upper dentition shows another combination of primitive and derived features, including a molariform P4, emerging parastyles and mesostyles, slender pericones and hypocones, and moderately developed buccal shearing crests that are mesiodistally straight. The morphology of Oligocene G. singsilai described by Marivaux et al. (2002) supported the suggested evolutionary trends of Sivaladapidae hypothesized by Qi and Beard (1998), demonstrating the enhancement of shearing crests and reduction (or complete absence) of both the pericone and the hypocone in the upper molars and that of the paraconid over time.

Furthermore, the transitional taxa connecting the more primitive Eocene Hoanghoniinae with the later Miocene sivaladapid species demonstrate that Hoanghoniinae is most likely a paraphyletic grouping from which Miocene sivaladapids are derived (Qi and Beard, 1998; Marivaux et al.,

2002).

15 While fossil discoveries and subsequent phylogenetic analyses in recent years have brought some resolution to our understanding of sivaladapid evolutionary relationships (Qi and

Beard, 1998; Ni et al., 2016; Gilbert et al., 2017), the question of sivaladapid origins and their phylogenetic affinities within Adapoidea remains unanswered. Previous studies have hinted at a close relationship to the European cercamoniid Periconodon (Qi and Beard, 1998; Gilbert et al.,

2017). Qi and Beard (1998) argue that Periconodon may be closely related to the sivaladapids based on the presence of a combination of derived dental features share between Pericondon,

Guangxilemur, Hoanghonius, and Rencunius. These four taxa possess a pericone and hypocone on the upper molars, a combination of features not seen in other adapoids. Results of a phylogenetic analysis performed by Qi and Beard (1998) also places Periconodon close to the basal sivaladapids.

A close evolutionary relationship between sivaladapids and asiadapids has also been proposed (Rose et al., 2009; Godinot, 2015). Asiadapiidae includes two genera and species,

Marcogodinotius indicus and Asiadapis cambayensis. Rose et al. (2009) suggested a potential ancestor-descendent relationship between asiadapids and sivaladapids based on a few derived dental features shared between the clades and their close biogeographic distribution, a hypothesis that proposes the asiadapids giving rise to the sivaladapids. In a synthesis describing the early

Eocene asiadapids from Vastan Mine in Gujarat, India, Rose et al. (2009) highlight the similarity between Marcgodinotius and Paukkaungia, despite Paukkaungia being approximately 50% larger in body size. The authors point out shared lower molar morphology including a simple P4, relatively transverse lower molar paracristids, and a deep trigonid (lingual) notch. Direct comparison of Marcgodinotius upper molars to that of Paukkaungia is currently impossible, given that upper molars are unknown for Paukkaungia; however, Rose et al. (2009) argue that

16 the morphology dP4 of Guangxilemur is comparable to that of Marcgodinotius, but is much larger. Furthermore, sivaladapid upper molars are very derived compared to other adapoids in their possession of both a hypocone and a pericone, so comparison of the upper molars of

Marcgodinotius to any sivaladapid at this point does not provide overwhelming support for a close relationship between asiadapids and sivaladapids.

Because asiadapids have been hypothesized to be closely related to sivaladapids, it is suitable to also note here the morphological similarities that have also been observed between the asiadapids and several European taxa (Rose et al., 2009; Godinot, 2015). In general, the asiadapids can be characterized as plesiomorphic, exhibiting dental morphology that most closely resembles Donrussellia (Bajpai, 2005; Rose et al., 2007; Rose et al., 2009; Godinot,

2015). Marcgodinotius is similar to Donrussellia in its overall shape and size, retention of four premolars, possessing a small, single-rooted lower P1, a two-rooted lower P2, and overall simple premolars (Rose et al., 2009). Features that distinguish Marcgodinotius from Donrussellia include more transverse lower molar protocristids, an indistinct or incipient paraconid on lower

M2-3, and a smaller hypoconulid lobe on lower M3 (Bajpai, 2005; Rose et al., 2009). These differences could be primitive and would therefore warrant a reappraisal of the adapoid ancestral morphotype (Godinot, 2015; Rose et al., 2009). A clear understanding of the polarity of certain dental characters would be required to firmly establish Marcgodinotius as primitive to

Donrussellia (Rose et al., 2009).

Asiadapis closely resembles Marcgodinotius in its dental morphology but differs in its slightly larger size and loss of P1 (Rose et al., 2009); while Asiadapis is derived in this manner, both Rose et al. (2009) and Godinot (2015) maintain the characterization of the species as primitive given the other plesiomorphic aspects of its cusp morphology. The upper dentition of

17 Asiadapis show primitive similarities to Cantius and Donrussellia, but slight differences distinguish Asiadapis as a distinct taxon. For example, the three taxa share a generalized, primitive cusp morphology and organization of P3, exhibiting a large paracone with a slightly smaller and lower protocone; however, Asiadapis differs in lacking a metastyle, which is present in Donrussellia (Rose et al., 2009). The lower teeth of Asiadapis also correspond closely in size and cusp morphology to that of Periconodon and Agerinia. The size and organization of crests of lower P4 in Periconodon are very similar to the condition seen in Asiadapis; Periconodon and

Asiadapis also share similar molar proportions and reduced paraconids. Lower molars of

Asiadapis are comparable to Agerinia in having an arcuate paracristid, extremely reduced (or indistinct) paraconids, and a deep trigonid notch (Rose et al., 2009).

Marcgodinotius also shows similarities to other cercamoniids, resembling Protoadapis,

Agerinia, and Periconodon in its plesiomorphic dental traits (Rose et al., 2009). These shared primitive traits include a single-rooted P1, a simple P4 with reduced or absent paraconid and metaconid, and overall reduced molar paraconids. Asiadapids differ slightly in their dental morphology from the cercamoniids, and those differences have been argued as evidence of a systematic distinction between the groups (Godinot, 2015; Rose et al., 2009). Traits distinguishing the asiadapids include a deep groove between the protoconid and metaconid in lower M2-3, and the preprotocrista directed toward a paraconule that continues toward the tip of the paracone in upper M1.

Taken in sum, asiadapids show great similarity in their dental morphology to European cercamoniids, and it could be argued that the asiadapids are closely related to the cercamoniids given the large number of shared features. Currently, there appears to be more evidence to suggest asiadapids could be more closely related to cercamoniids than to sivaladapids; although

18 Rose et al. (2009) and Godinot (2015) propose a close relationship between sivaladapids and asiadapids, the evidence to support the relationship is not as strong as the evidence supporting an asiadapid-cercamoniid connection. Recent phylogenetic analyses have offered conflicting results regarding the phylogenetic affinities of asiadapids. For example, results from the phylogenetic analysis performed by Rose et al. (2009) place both Asiadapis and Marcgodinotius the base of a clade that includes in a successive branching pattern the hoanghoniines + Wailekia, and

Agerinia, which does suggest a close asiadapid-sivaladapid connection. Other phylogenetic analyses have recovered clades showing a close relationship between the asiadapids and

Donrussellia (Seiffert et al., 2009; Ni et al., 2016; Gilbert et al., 2017; Seiffert et al., 2018).

These results still leave the question of the phylogenetic affinities of sivaladapids unanswered, but they highlight a possible broader Eurasian relationship rather than an explicit asiadapid- sivaladapid connection; if the asiadapids are more similar in their morphology to cercamoniids, and sivaladapids have been described as having similar morphology to Periconodon, then a close relationship between the sivaladapids to a European clade falls logically under this evolutionary narrative and biogeographic pattern. The larger number of similarities between the asiadapids and the cercamoniids further questions the hypothesis of a close or direct relationship between the asiadapids and the sivaladapids.

Thus, the goal of this study is to look more closely at the origins of the Sivaladapidae and test these competing hypotheses about the origins of these late-surviving Asian taxa. To answer these questions, this study takes a slightly different approach to conducting a phylogenetic analysis than has been tried before with this group. Here, only agreed-upon basal sivaladapid taxa (e.g., hoanghoniines) were compared to other basal adapoids representing different adapoid families with respective biogeographic distributions spanning North America, Europe, and Asia

19 (see Figures: 5-8; see Table 1). The objective of comparing only the basal taxa from different adapoid families is to observe any distinct synapomorphies uniting sivaladapids with another adapoid group before the evolution of autapomorphies in the various respective adapoid families.

This strategy thus helps to prevent the chance of long branch attraction in phylogenetic analysis, a general phenomenon that can lead to inaccurate phylogenetic hypotheses (Bergsten, 2005).

Materials and Methods

Fossil specimens examined in this study derive from several different collections (see

Tables: 3-14). Morphometric data were taken largely from the published literature. When certain measurements were not available in the literature, published photographs of the fossil specimens were uploaded into the image processing software ImageJ (1.51s) which allows users to obtain metric measurements from images. To capture non-traditional measurements in

ImageJ, pixels of uploaded fossil images were converted to metric units based on known distances, creating a metric scale by which dimensions were then compared and recorded. This method allowed for previously unpublished data to be incorporated into the analyses. It also allowed for the addition of new, novel data (i.e., non-traditional morphometric data/measurements) to be supplemented to the traditional dimensions in published in descriptions of fossil specimens. All dental measurements were recorded following landmarks and descriptions given by Gilbert et al. (2017).

In addition to the more traditional morphometric data that were recorded from the dentition, a number of comparative relative measurements were recorded following Gilbert et al.

(2017) (see Tables: 4, 6, 9, 10). These indices compare relative dental sizes and shapes; they also incorporate the non-traditional measurements to quantitatively compare the size and shape of regions within a single tooth. In other words, the comparative relative methods compare

20 intraspecific and interspecific variation in dental size and shape. Raw data was recorded for the taxa sampled and population averages were then calculated. The population averages served as the value by which to score each taxon for a given character for the phylogenetic analysis (see additional comments on scoring method).

To understand the origin of Sivaladapidae, a phylogenetic analysis containing dental and mandibular characters was performed. The analysis included a total of 82 qualitative and quantitative characters, largely building on analyses of Gilbert et al. (2017) and Qi and Beard

(1998) (Table 2). Additional characters were also derived from Ni et al. (2016) and independent observations. For taxa represented by an M1, body mass was estimated using the M1 area prosimian regression equation developed by Conroy (1987) or taken from Gilbert et al. (2017), who also used this equation. As outlined in Gilbert et al. (2017), taxa not preserving a lower M1

(e.g., Adapoides) were compared with other taxa with similar upper molar area dimensions. For example, Adapoides and Donrussellia differ only slightly in M2 area. Thus, body mass for

Adapoides was estimated to be similar to that of Donrussellia (the close comparison of these taxa is further supported in the literature, see Gunnell et al., 2008). With the exception of three characters describing the orientation of the cristid obliqua, all 82 characters were ordered and equally weighted. For quantitative characters, the gap weighted coding method (Thiele, 1993) was applied, using on average three character states, which allows for the range of data in a given sample to dictate character state (i.e., score) designations based on the gaps in between values; when comparing size and shape of specimens, this method is preferred over qualitative character scoring as it incorporates the morphometric data in an objective manner to assign scores rather than relying on subjective score assignment, which could differ by observer.

21 In order to account for as many factors as possible, gain overall better representation of fossil taxa in the analysis, and evaluate the overall robusticity of the resulting trees, two different sets of analyses were performed. One set can be identified as a “conservative” analysis, while the other is identified as a “liberal” analysis. Both analyses use the same tree searching parameters but differ slightly in their morphometric data representation (see Tables: 11-14). The data matrix comprising the conservative analysis did not include fossils identified in the literature as either M1/2, but rather included only those definitively assigned to a specific molar position. Additionally, in the conservative analysis comparative-relative measurements (indices) were not calculated at the population level; in other words, all comparative-relative measurements were derived from individuals (i.e., specimen) that preserved both elements being compared (Tables: 11, 13). Data compilation of this manner has its merits, in that it represents observed comparative-relative indices that we can be confident in, as they derive from specimens with accurately identified teeth which then can be measured in the calculation of these indices.

However, this manner of data compilation also significantly restricts the quantity of data to analyze. Sample sizes for comparisons using this method were low and, in some cases, could not be calculated given the “conservative” method parameters (i.e., that the two dimensions being compared must come from the same individual). To account for low representation (a reality of the fossil record), and to include as much data as possible, the liberal analysis incorporated specimens identified as either M1/2 (assignment to molar position, for example, was assigned on a case by case basis for each specimen). The liberal analysis also took comparative relative measurements at the population level for each taxon, not only creating more data, but also better represented data (Tables: 12, 14). Taking these indices at the population level should be, in theory, close to the average of indices from individual specimens. Preparing conservative and

22 liberal morphological datasets using the parameters described above permitted the observation of what the most conservative value for a comparative-relative measurement would be for a given taxon which could then be compared to the calculated population averages from a larger sample size that took into consideration more specimens. It could be argued that preparing the data in this manner could help evaluate particular specimens and dimorphism in a species. In the case of the taxa sampled in the current analysis, population averages were not significantly altered with the addition of more specimens.

The resulting matrix of 82 qualitative and quantitative characters, for both conservative and liberal datasets, was analyzed using a random addition sequence Tree-Bisection-

Reconnection search, in which 10,000 replications were run to find the most parsimonious tree

(MPT) in the parsimony analysis program TNT, Tree analysis using New Technology (Goloboff et al., 2008). Clade support estimates were generated by running a 1,000 replication bootstrap procedure. Bremer support values, retention indices, and consistency indices were also generated to evaluate clade support. Strict and majority-rule consensus trees of all trees within

50% of the MPT were also constructed for all analyses, respectively. Only basal adapoid taxa representing five different adapoid families were examined as part of the ingroup while

Purgatorius, Altanius, and Teilhardina were assigned and constrained sequentially as the outgroup. Purgatorius and Teilhardina were represented by the earliest known species (i.e., P. titusi, P. unio, T. asiatica, T. magnoliana, T. belgica) assigned to the genus, respectively. These taxa represent the earliest occurring species of the genera and, in most cases, exhibit overall primitive morphology compared to later occurring Purgatorius and Teilhardnia species.

Composing the ingroup of only basal adapoids was fundamental in addressing sivaladapid origins for this analysis. All trees listed here (Images 9-12) were produced using FigTree v1.4.3.

23 The liberal and conservative analyses differ slightly in their data compilation as mentioned above, but also by the manner in which certain taxa were represented. In the conservative analysis, Cantius was represented by two species, C. ralstoni and C. torresi. For the liberal analysis, these two species were combined and data for both species were represented at the genus level. In total, four independent tree searches were performed: two searches each for the liberal and conservative datasets. The difference in these searches are related to representation and then removal of three sivaladapid taxa, Guangxilemur tongi, Guangxilemur singsali, and Wailekia orientale. Initial inclusion of these taxa was based on their more basal placement within Sivaladapidae than the later Miocene forms (Qi and Beard, 1998; Gingerich et al., 1994). Removal of these taxa for subsequent tree searches was justified due to their intermediate and more derived morphology compared to the earlier hoanghoniines. To avoid skewing the analysis given the more derived morphology of Guangxilemur and Wailekia, these taxa were removed in later tree searches.

Results

Results for both sets of analyses are not well resolved (Figures 9-12). For the conservative datasets, two independent parsimony analyses were performed; the first search included taxa representing Guangxilemur and Wailekia and examined 22 taxa total, while the second search excluded these taxa and examined 19 taxa total. The two analyses recovered 690 and 10 most parsimonious trees (MPTs) with lengths of 216 and 196, respectively (Figure 9;

Figure 11). The strict consensus tree recovered from examination of 19 taxa is more resolved than that of the analyses that examined 22 taxa. Removal of Guangxilemur and Wailekia from the ingroup led to an increase in overall phylogenetic resolution, suggesting that the poor representation of Guangxilemur could have affected the degree of resolution. Both conservative

24 analyses recovered clades in which the European adapoid taxa are placed among the Asian taxa

(Figures: 9, 11); in the analysis that excluded Guangxilemur and Wailekia, the majority-rule consensus found a more deeply nested position of Periconodon, placing it as the sister taxon to a clade that includes (Agerinia (Hoanghonius + Rencunius)) (Figure 11). Neither analysis examining the conservative datasets recovered a sister-taxon relationship of Paukkaungia +

Kyitchaungia, which is interesting given that the analyses performed by Gilbert et al. (2017) repeatedly recovered this particular grouping. Marcgodinotius and Asaiadpis are placed outside a paraphyletic Sivaladapidae. Bootstrap support values and Bremer support values for both conservative analyses are extremely low; no unconstrained clades are supported over 50% and recovered decay indices are one.

For the liberal datasets, two independent parsimony analyses were also performed following the protocol regarding the representation of Guangxilemur and Wailekia described above. The first search examined a total of 21 taxa and recovered 31 MPTs with a length of 221

(Figure 10). The second search examined a total of 18 taxa and recovered only one MPT with a length of 200 (Figure 12). The strict consensus tree recovered in the analysis that examined only

18 taxa is again much more resolved. In both searches, Hoanghonius and Rencunius are supported as sister taxa. A clade containing (Europolemur (Protoadapis + Lushis)) is recovered as the sister group to a clade containing (Periconodon (Agerinia (Rencunius + Hoanghonius))).

Support for both analyses are also low; Hoanghonius + Rencunius is the only clade with a bootstrap support value over 50% (Figure 12) and every node has a decay index of one.

Given the overwhelming evidence indicating that the removal of Guangxilemur and

Wailekia produced an overall increase in phylogenetic resolution in both conservative and liberal analyses, it is reasonable to compare from this point forward only the MPTs recovered in the

25 analyses that eliminated these taxa from the ingroup. Both analyses that examined conservative and liberal datasets share points of similarity in their consensus trees. For example, both majority-rule consensus trees recover a successive branching pattern of (Periconodon (Agerinia,

(Rencunius + Hoanghonius))); this clade differs in its placement though, with the conservative analysis placing this clade at an earlier branching position than the liberal analysis. They also recover a sister-taxa relationship between Lushius and Protoadapis. Their points of difference are also interesting to note. For example, the conservative analysis places Donrussellia as the most primitive taxon just outside of the group of branching relationships that include the

Eurasian adapoids and sivaladapids. In the liberal analyses (Figures: 10, 12), however, Panobius is placed in an earlier branching position just before Donrussellia, both of which are nested outside the group containing the remaining Eurasian taxa and sivaladapids. In short, Panobius has a deeply nested position in the conservative analyses (Figures: 9, 11) but jumps to a much more primitive position in the liberal analysis. The varying in position of Panobius could be related to the incorporation of more premolar character data in the liberal analysis; Gunnell et al.

(2008) discuss the overall primitive morphology of Panobius, suggesting that the taxon may appear in a more primitive phylogenetic position depending on the polarity of premolar characters. Compared to previous studies that have described Marcgodinotius and Asiadapis as extremely primitive in their dental morphology (Rose et al., 2009; Godinot, 2015), all four independent analyses here place these taxa at the base of a clade that branches off after

Donrussellia or even more deeply nested within that clade. These two asiadapid taxa are also repeatedly found in close connection to Adapoides and Kyitchaungia.

In short, the phylogenetic hypotheses favored here are those that exclude Guangxilemur and Wailekia, with further preference of the majority-rule consensus tree produced by examining

26 the liberal dataset (i.e., Figure 12). Taken in sum, the four independent searches produced MPTs that recovered similar relationships, most notably the affinity of sivaladapids with Eurasian adapoid taxa rather than the North American notharctids.

Discussion

Sivaladapids have been known to paleontologists for decades, yet their origins and phylogenetic affinities to other adapoid groups have remained enigmatic. Fossil prospecting of

Asian localities spanning the early Eocene to the late Miocene has produced a more species rich

Sivaladapidae, with identification of a number of new genera and species over the past 25 years

(Gingerich et al., 1994; Qi and Beard, 1998; Beard et al., 2007; Ni et al., 2016; Gilbert et al.,

2017). Only recently have phylogenetic relationships within Sivaladapidae been more thoroughly investigated, finding some resolution among the later Miocene sivaladapines, and newly named subfamilies (Qi and Beard, 1998; Ni et al., 2016; Gilbert et al., 2017). Previous studies have also aimed to place some sivaladapid taxa within the larger primate radiation (Ni et al., 2016). While studies of sivaladapids and adapoids broadly have traditionally questioned their potential connection or ancestral relationship to extant strepsirrhines and haplorhines, the current study focuses on the phylogenetic affinities of Sivaladapidae relative to other adapoid families. This study broadens the lens of phylogenetic inquiry of sivaladapids outside the family level to understand the origin of sivaladapids in terms of which adapoid family they share a close evolutionary relationship by performing a phylogenetic analysis examining basal adapoid and sivaladapid taxa.

Results of the four independent parsimony analyses recovered MPTs with overall low phylogenetic resolution, which is to be expected when that dataset is limited by fragmentary fossil evidence and therefore contains a lot of missing data. Additionally, when compared to

27 other studies (Ni et al., 2016; Seiffert et al., 2005), the dataset analyzed here can be characterized as limited, given the small number of dental characters. Low resolution could also be related to the taxa sampled in the analysis; while sampling only basal adapoid taxa prevents the potential problem of long branch attraction, it runs the risk of examining a sample in which no phylogenetic signals exist. In other words, the taxa that are close to the stem may have experienced a fast and bushy radiation, thus leaving little differentiation or phylogenetically significant differences in the morphology. Furthermore, it has been previously demonstrated that increasing the number of taxa and characters in an analysis generally increases accuracy, and therefore may outweigh any potential long-branch attraction (Wiens, 2003, 2005; Heath et al.,

2008). Nevertheless, the current results offer new information worthy of discussion.

The strict-consensus and majority-rule trees from the four analyses vary in their topologies but certain clades were repeatedly recovered among all trees, suggesting some support for these groupings. The preferred majority-rule consensus tree here (from the analysis that examined 18 taxa from the liberal dataset) recovered two clades demonstrating a close relationship between the sivaladapids and Eurasian adapoids. Repeatedly recovered groupings include a sister-taxa relationship between Hoanghonius and Rencunius that is nested within a clade containing Periconodon followed by Agerinia, and a sister taxon relationship between

Lushius and Protoadapis. A relationship between Periconodon and Eocene sivaladapids has been previously hypothesized based on their shared morphology of possessing both a hypocone and pericone on the upper molars and the presence of molar conules (Qi and Beard, 1998). The recovered Lushius + Protoadapis clade in both liberal and conservative analyses is interesting, given that Lushius is very poorly known and considered an enigmatic sivaladapid in itself with uncertain phylogenetic affinity within Sivaladapidae. The M2 of these taxa are extremely similar

28 in overall dimensions (size and shape). The repeated recovery of these European and sivaladapid clades suggests that the classic hoanghoniines, Hoanghonius and Rencunius, and perhaps

Lushius are connected with European taxa, particularly the cercamoniids. Meanwhile, the more recently discovered and fragmentary Paukkaungia and Kyitchaungia may be more closely related to asiadapids and caenopithecids than to other sivaladapid taxa, as indicated by the analyses excluding Guangxilemur and Wailekia, suggesting that Sivaladapidae as currently constituted may be paraphyletic. More broadly, the results of these analyses do not support a relationship between any sivaladapids and notharctids, and instead suggest a complex phylogenetic and evolutionary scenario among Eurasian adapoids.

Gunnell et al. (2008) discuss the paleobiogeographic patterns among the northern continents during the onset of the Paleocene-Eocene Thermal Maximum, describing two routes connecting Asia with Europe and North America as the Indo-Pakaistan subcontinent collided with Asia. These routes are highlighted by apparent faunal interchange patterns by which the

Early Eocene primates likely migrated between Asia and Europe. This European-Asian faunal exchange pattern is supported by a hypothesized close relationship between Donrussellia and

Panobius, a relationship that is also recovered in the current study’s preferred majority-rule consensus tree; it also reinforces a close relationship between the Asian sivaladapids and the

European taxa. While Gunnell et al. (2008) also discuss a passageway from Asia to North

America, east of the nascent Himalayas, the current analysis does not support the relationship between sivaladapids and notharctids.

Related to the discussion above, Beard et al. (2007) discuss the morphology of the partial skeleton NMMP 20 recovered from the Eocene Pondaung Formation of Myanmar that had been previously identified (and fervently debated) as an amphipithecid and potentially having

29 anthropoid affinities (Ciochon et al., 2001). Beard et al. (2007) argue that the postcrania of

NMMP 20 more closely resembles the calcaneus of the sivaladapid Kyitchaungia, and thus identify NMMP 20 as a sivaladapid with possible connection to Guangxilemur due to its large body size. However, Ciochon et al. (2001) also describe the postcranial morphology present in

NMMP 20 that resembles notharctids and maintain that the phylogenetic affinities of the fossil are unclear. If NMMP 20 is in fact a sivaladapid and not an amphipithecid as Beard et al.,

(2007) argue, then its postcranium is notharctid-like. The results from the current analysis, however, do not support a notharctid-sivaladapid relationship, and sivaladapids are recovered in all MPTs in deeply nested positions within clades completely separate from the sampled notharctid taxa (i.e., Cantius).

Another interesting pattern observed in the topologies of all majority-rule consensus trees is the placement of the asiadapid taxa, Marcgodinotius and Asiadapis. As mentioned above,

Rose et al. (2009) and Godinot (2015) suggested a close evolutionary relationship between the sivaladapids and asiadapids based on shared dental morphology, a hypothesis that was supported by one phylogenetic analysis performed by Rose et al. (2009). However, similar to other recent phylogenetic analyses (Seiffert et al., 2009; Ni et al., 2016; Gilbert et al., 2017; Seiffert et al.,

2018), this study does not find a close relationship between the asiadapids and a monophyletic

Sivaladapidae. The four independent parsimony analyses performed in this study produced varying tree topologies, shifting the asiadapids to different branching positions and placements.

Despite these varied positions, Marcgodinotius and Asiadapis are recovered most often in a grouping with Adapoides that is nested within a broader Eurasian clade. Dental comparisons between Asiadapis and Adapoides show similar lower molar morphology, including an elongated talonid of the lower M3 and a lingually placed (but also reduced) paraconid. The phylogenetic

30 analysis performed by Gilbert et al. (2017) had assigned Marcgodinotius to the outgroup and produced results showing a close relationship between the sivaladapids and Pericondon; the authors note, however, that a different relationship could be recovered if Marcgodinotius were not assigned and constrained to the outgroup. The current analysis assigned Marcgodinotius to the ingroup, and a direct relationship between the asiadapids and sivaladapids was still not found, but connections between Asian and European taxa were repeatedly recovered. Taken in sum, the current analysis does not find substantial support for a close or direct relationship between the asiadapids and Sivaladapidae, which contrasts with the views of Rose et al. (2009) and Godinot

(2015). However, if the analyses excluding Guangxilemur and Wailekia are more accurate, it is possible that asiadapids are closely related to some taxa currently recognized as sivaladapids, namely Paukkaungia and Kyitchaungia. In either case, what this study and previous studies

(Seiffert et al., 2018, 2009; Gilbert et al., 2017; Ni et al., 2016) have pointed out is that there appears to be some kind of close relationship and complex biogeographic interchange between the Asian and European adapoid taxa. The shared morphology noted between the asiadapids and the European taxa could be taken as support for a sivaladapid and European taxa connection, given the overall pattern of European taxa grouping.

Also worth repeating from the current study is the nature of Sivaladapidae in the results of all four independent parsimony analyses. Each analysis recovers a paraphyletic

Sivaladapidae, in that sivaladapid taxa are recovered in different clades with different European adapoid taxa. These results may speak to the instability of the family and question the integrity of some of the synapomorphies linking all sivaladapid taxa. Given the repeatedly recovered connections to European taxa, it is possible that some of the previously identified sivaladapid synapomorphies may not actually be phylogenetically significant (i.e., twinned hypoconid and

31 entoconid). Perhaps other dental features that link sivaladapids to the European taxa (i.e., premolar/molar size and shape) are better/more accurate phylogenetic signals. For example,

Gilbert et al. (2017) found a sister taxa relationship between Paukkaungia and Kyitchaungi, and the twinned hypoconulid and entoconid feature was interpreted in part as being one of the synapomorphies closely uniting the two taxa as sivaladapids; however, the current study does not support this relationship. Building on the character matrix from the Gilbert et al. (2017) analysis, the current analysis introduced additional qualitative characters and premolar size and shape characters. When the new characters were incorporated, along with other taxa assigned to the ingroup, the close sister taxa relationship between Paukkaungia and Kyitchaungia was no longer found. This result may also simply be a product of the innate nature of phylogenetic investigation and parsimony analysis, in that new characters and new taxa may recover different relationships, but it does question the integrity of some the synapomorphies of sivaladapids as actual phylogenetic signals.

Conclusion

Advances in our understanding of early primate evolution have been facilitated by continuous investigation of the fossil record and subsequent phylogenetic analyses, which in turn have produced several competing hypotheses of the phylogenetic affinities of different fossil primate clades (Hofstetter, 1977; Gingerich and Simon, 1977; Gingerich, 1977a, 1977b; Kay and

Cartmill, 1978; Rasmussen, 1986; Seiffert at el., 2009; Gingerich, 2012; Ni et al., 2013).

Focusing on one extinct primate clade in particular, this study contributes new information to the ongoing investigation and discussion of sivaladapid origins and their phylogenetic affinities within Adapoidea. The phylogenetic analysis performed here sampled sivaladapids and other basal adapoid taxa in a novel way to test the previously proposed hypotheses of sivaladapid

32 phylogenetic affinities (Rose et al., 2009; Godinot, 2015). Broadly, results from the current analysis suggest that sivaladapids share a closer evolutionary relationship with European adapoids (i.e., cercamoniids), rather than a close relationship with North American taxa (i.e., notharctids). These results contradict the hypotheses of both Rose et al. (2009) and Godinot

(2015), at least in part. In order to resolve the ongoing debate of sivaladapid evolutionary relationships, an analysis incorporating postcranial data will be necessary, as additional anatomical representation (particularly postcrania) and new data in the form of new fossil material may enhance the investigative quality of the analysis. Additionally, performing another type of analysis (i.e., Bayesian) might offer another interesting perspective. In sum, additional specimens, characters, and analytical perspectives will be necessary to more confidently flesh out the evolutionary relationships of sivaladapids within Adapoidea.

Acknowledgements

I would like to express my deep gratitude for the mentorship and guidance offered by my thesis advisor, Dr. Chris Gilbert; he has fostered my academic development as a student and instilled a passion for the study of early primate evolution. His support and direction during the entirety of this project was invaluable and greatly appreciated. I also acknowledge and thank Dr.

Stephen Chester for serving as my second reader, whose comments and feedback improved this study. I thank Kelsey Pugh for her endless support and assistance in the methodological aspects of this project. I would also like to recognize my friends and colleagues at the AMNH for their words of encouragement and advice during the many stages of this study. I extend immense gratitude toward Dr. Julia Zichello, who has also served as a colleague and mentor; I am deeply thankful for her insights and advice on this project. I thank my parents, Dr. Stace Rust and

Jeffrey Rust, for their unwavering support. I thank my grandmother, Elaine Staglik, who always

33 reminded me to consider the relevance, value, and importance of this study. Lastly, I thank

Stephen Venner for always believing in me.

Figures and Tables

Figure 1: Upper dental morphology typifying hoanghoniines. a) University of Uppsala (no number), right M2 of Hoanghonius stehlini, and b) IVPP 5311, right P4-M1 of Rencunius zhoui, in occlusal views. Red arrows point to the molar pericone and hypocones that arise from a lingual cingulum, a feature that characterizes hoanghoniines. P4 of hoanghoniines are simple and not molarized compared to later Miocene sivaladapids, lacking conules and accessory cusps. Image modified from Gingerich et al. (1994).

34

Figure 2: Lower dental morphology typifying hoanghoniines. a-b) Type specimen of Hoanghonius stehlini, University of Uppsala (no number,) left dentary preserving M2-3 in occlusal and lateral views and, c-d) Type specimen of Rencunius zhoui, IVPP 5312, left dentary preserving P4-M2 in occlusal and lateral views. Yellow arrows point to twinned entoconid and hypoconulid that are situated lingually, a feature that unites all sivaladapid taxa. The relatively low-crowned molars and simple, unmolarized P4 distinguish the hoanghoniines from later sivaladapids that possess highly-crested molars and molarized P4. Hoanghoniine trigonids are not as mesiodistally compressed compared to sivaladapines. Image modified from Gingerich et al. (1994).

35

Figure 3: Upper and lower dentition of Guangxilemer. a) Holotype of G. tongi, IVPP V11652, left M2 in occlusal view; b) Holotype of G. singsila, DBC 2168, right M2 in occlusal view; c) DBC 2165, right dP4 of G. singsilai in occlusal view; d) DBC 2170, right M1/2 of G. singsilai in occlusal view. Guangxilemur displays intermediate dental morphology, exhibiting features that are transitional between the hoanghoniines and sivaladapines. Red arrows point to pericones and hypocones of the upper molars, which are present but reduced in G. singsilai compared to the hoanghoniines; pericones and hypocones are secondarily lost in sivaladapines. Blue arrows point to the stylar cusps (i.e., parastyle and mesostyle) that enhance the external shearing crests of the upper molars, producing a W-shaped structure that also typifies later sivaladapines. Yellow arrows point to the twinned hypoconulid and entoconid. Images modified from Qi and Beard (1998) and Marivaux et al. (2002).

36 Figure 4: Upper and lower dentition Sivaldapis nagrii. a) LUVP 14506, left maxilla 3 1 preserving P -M in occlusal view and, b-c) UM 71837, left dentary preserving P4-M1 in lateral and occlusal views. Red arrows point to the well-developed upper molar parastyles 4 and mesostyles, which act to enhance upper molar shearing crests. Note the molarized P in figure 4a, which is a unique feature evolved in the Miocene sivaladapines. White arrows point to the strong upper molar lingual cingulum, a typical feature found in Miocene sivaladapines. Yellow arrows point to the twinned hypoconulid and entoconid found in all sivaladapids. Sivaladapines exhibit lower molars trigonids that are mesiodistally compressed compared to hoanghoniines. Images modified from Gingerich and Sahni (1984).

37 Table 1: Taxa sampled in current study Geographic Estimated Taxon (Family; Genus & Species) Geologic Age Distribution Body Mass (g)

Incertae sedis Altanius orlovi Early Eocene Asia 21 Purgatoriidae Purgatorius spp. Early Paleocene North America 103 Asia, Europe, Omomyidae Teilhardina spp. Early Eocene North America 69 Notharctidae Cantius torresi Early Eocene North America 558 Notharctidae Cantius ralstoni Early Eocene North America 814 North America, Notharctidae Cantius spp. Early Eocene Europe* 656 Cercamoniidae Donrussellia provincialis Early Eocene Europe 133 Middle to Late Caenopithecidae Europolemur klatti Eocene Europe 771 Cercamoniidae Periconodon huerzeleri Middle Eocene Europe 331 Cercamoniidae Agerinia roselli Middle Eocene Europe, Asia* 419 Early to Middle Cercamoniidae Protoadapis curvicuspidens Eocene Europe 1207 Asiadapidae Marcgodinotius indicus Early Eocene Asia 109 Asiadapidae Asiadapis cambayensis Early Eocene Asia 273 Caenopithecidae Adapoides troglodytes Middle Eocene Asia 133 Cercamoniidae Panobius russelli Late early Eocene Asia 86 Sivaladapidae Hoanghonius stehlini Middle Eocene Asia 684 Sivaladapidae Rencunius zhoui Late middle Eocene Asia 733 Sivaladapidae Lushius quilinensis Late Eocene Asia 1450 Sivaladapidae Wailekia orientale Late Eocene Asia 1012 Sivaladapidae Paukkaungia parva Late middle Eocene Asia 483 Sivaladapidae Kyitchaungia takaii Late middle Eocene Asia 1040 Sivaladapidae Guangxilemur tongi Late Eocene Asia 4800 Sivaladapidae Guangxilemur singsilai Early Oligocene Asia 1526 Body mass estimates represented here derived from M1 area prosimian regression equation developed by Conroy (1987) or taken from Gilbert et al. (2017). *Indicates genus has wider biogeographic distribution than particular species sampled in this analysis.

38

Figure 5: Map showing the biogeographic distribution of all fossil taxa sampled in the analyses (Google Maps, 2018)

39

Figure 6: Map showing the biogeographic distribution of all North American fossil taxa sampled in the analyses (Google Maps, 2018). Species representing Purgatroius here include P. unio and P. titusi. Species representing Teilhardina here include T. magnoliana.

40

Figure 7: Map showing the biogeographic distribution of all European fossil taxa sampled in the analyses (Google Maps, 2018). Species representing Teilhardina here include T. belgica. Refer to Table 1 for species level of remaining taxa represented here.

41 Figure 8: Map showing the biogeographic distribution of all Asian fossil taxa sampled in the analyses (Google Maps, 2018). Species representing Teilhardina here include T. asiatica.

Refer to Table 1 for species level of remaining taxa represented here.

42 Table 2: Characters used in phylogenetic analysis Character Definition and character states Type Reference Lower molar 1 0 = absent/weak, 1 = present and strong QL Qi and Beard (1998) hypoconulid shape lower molar 2 0 = buccal, 1 = central, 2 = twinned w/ entoconid QL Qi and Beard (1998) hypoconulid position Lower molar lingual 3 0 = absent/weak, 1 = present and strong QL Qi and Beard (1998) notch M paraconid Gilbert et al. (2017); 4 1-2 0 = absent/indistinct, 1 = polymorphic, 2 = present/distinct QL development Ni et al. (2016) 0 = distal to the protoconid, 1 = near midline between protoconid and metaconid, 2 = M cristid obliqua Gilbert et al. (2017); 5 1 connected to the metaconid, 3 = connected to protoconid near buccal border, 4 = QL, U orientation Ni et al. (2016) polymorphic between states 0 and 1 M cristid obliqua 1 = distal to the protoconid, 1 = near midline between protoconid and metaconid, 2 = Gilbert et al. (2017); 6 2 QL, U orientation connected to the metaconid, 3 = connected to protoconid near buccal border Ni et al. (2016) M cristid obliqua 2 = distal to the protoconid, 1 = near midline between protoconid and metaconid, 2 = Gilbert et al. (2017); 7 3 QL, U orientation connected to the metaconid, 3 = connected to protoconid near buccal border Ni et al. (2016) Entoconid-hypoconulid 8 0 = absent/weak, 1 = present and strong QL Gilbert et al. (2017) notch Lower molar crown 9 0 = moderate, 1 = high-crowned QL Qi & Beard (1998) height

10 P4 length relative to M1 0 = short, 1 = long QN Gilbert et al. (2017) Upper and lower P4 11 0 = premolariform, 1 = submolariform, 2 = molariform QL Qi & Beard (1998) molarization

12 P4 shape 0 = short/broad, 1 = long/narrow QN Gilbert et al. (2017) P paraconid Gilbert et al. (2017); 13 4 0 = abset/weak, 1 = present/strong QL development Ni et al. (2016) P talonid cusp 0 = one distinct cusp present, 1 = two distinct cusps present, 2 = polymorphic between two Gilbert et al. (2017); 14 4 QL development and three cusps present, 3 = three distinct cusps present Ni et al. (2016) 15 M1-2 hypocone 0 = abset, 1 = present and weak/small, 2 = present and strong/large QL Qi & Beard (1998)

16 M1-2 preicone 0 = absent, 1 = present and weak/small, 2 = present and strong/large QL Qi & Beard (1998) Lingual cingulum on 17 0 = weak/incomplete, 1 = strong/continuous QL Qi & Beard (1998) upper molars

43 18 M1-2 shape 0 = square, 1 = intermediate, 2 = markedly transverse QN Qi & Beard (1998)

19 M3 shape 1 = square, 1 = intermediate, 2 = markedly transverse QN Gilbert et al. (2017)

20 M1-2 molar conules 0 = distinct, 1 = indistinct or absent QL Qi & Beard (1998)

21 M1-2 parastyle 0 = weak/absent, 1 = strong QL Qi & Beard (1998)

22 M1-2 mesostyle 1 = weak/absent, 1 = strong QL Qi & Beard (1998)

M1-2 external shearing 23 0 = mesiodistally straight, 1= moderately W-shaped, 2 = W-shaped QL Qi & Beard (1998) crest shape

24 M1 area/upper M2 area 0 = M2 relatively large, 1 = M1 relatively large QN Gilbert et al. (2017)

25 M1 area/upper M3 area 0 = M3 relatively large, 1 = intermediate, 2 = M1 relatively large QN Gilbert et al. (2017)

26 M2 area/upper M3 area 1 = M3 relatively large, 1 = intermediate, 2 = M2 relatively large QN Gilbert et al. (2017)

27 Premolar number 0 = four, 1 = polymorphic four or three, 2 = three, 3 = two QL Gilbert et al. (2017)

28 P2 root number 0 = two, 1 = one QL Gilbert et al. (2017)

29 M1-2 shape 0 = short and broad, 1 = intermediate, 2 = long and narrow QN Gilbert et al. (2017) M trigonid 30 1-2 0 = mesiodistally compressed, 1 = uncompressed QN Gilbert et al. (2017) compression M trigonid lingual Gilbert et al. (2017); 31 1-2 0 = lingually open, 1 = polymorphic, 2 = lingually closed QL shape Ni et al. (2016) Lower molar buccal 32 0 = absent/weak, 1 = polymorphic/intermediate, 2 = strong/continuous QL Gilbert et al. (2017) cingulid strength

34 M3 shape 0 = short and broad, 1 = long and narrow QN Gilbert et al. (2017)

35 M3 trigonid shape 0 = compressed, 1 = intermediate, 2 = uncompressed QN Gilbert et al. (2017)

36 M1 area/lower M2 area 0 = M2 relatively large, 1 = intermediate, 2 = M1 relatively large QN Gilbert et al. (2017)

37 M1 area/lower M3 area 0 = M3 relatively large, 1 = intermediate, 2 = M1 relatively large QN Gilbert et al. (2017)

44 38 M2 area/lower M3 area 0 = M3 relatively large, 1 = intermediate, 2 = M2 relatively large QN Gilbert et al. (2017) Mandibular height at 38 0 = shallow, 1 = deep QN Gilbert et al. (2017) lower M1 39 Symphyseal fusion 0 = mandibular symphysis unfused, 1 = partially fused, 2 = fused QL Gilbert et al. (2017)

40 Estimated body mass 0 = small, 1 = intermediate, 2 = large QN Gilbert et al. (2017)

41 P1 evolutionary loss 0 = absent, 1 = present QL Ni et al. (2016)

42 P1 size relative to P2 0 = small P1, 1 = P1 slightly smaller to slightly larger QN Ni et al. (2016)*

43 P2 evolutionary loss 0 = absent, 1 = present QL Ni et al. (2016)

44 P2 size relative to P3 0 = much shorter, 1 = shorter, 2 = slightly shorter to similar in size QN Ni et al. (2016)* P crown height relative 45 2 0 = much shorter, 1 = shorter, 2 = slightly shorter to similar in size QN Ni et al. (2016)* to P3

46 P2 caniniform 0 = absent, 1 = present QL Ni et al. (2016)

P2 sectorial premolar - 47 development of honing 0 = absent or very weak, 1 = present QL Ni et al. (2016) facet

48 P2 shape 0 = short/broad, 1 = long/narrow QN Gilbert et al. (2017) P talonid cusp 0 = one distinct cusp present, 1 = two distinct cusps present, 2 = polymorphic between two Gilbert et al. (2017); 49 2 QL development and three cusps present, 3 = three distinct cusps present Ni et al. (2016)

50 P3 evolutionary loss 0 = absent, 1 = present QL Ni et al. (2016)

51 P3 length relative to P4 0=much shorter, 1=shorter, 2=slightly shorter to similar in size QN Ni et al. (2016)*

52 P3 length relative to M1 0=much shorter, 1=shorter, 2=slightly shorter to similar in size QN Ni et al. (2016)*

P protoconid height 53 3 0=much shorter, 1=shorter, 2=slightly shorter to similar in height QN Ni et al. (2016)* relative to that of P4

Lower P3 protoconid 54 height relative to that of 1 = P3 lower, 1 = similar in height, 2 = P3 higher QN Ni et al. (2016) M1

45 P paraconid 55 3 0 = absent/weak, 1 = present/strong QL Ni et al. (2016) development

56 P3 metaconid size 0 = absent, 1 = small distinct cusp, 2 = large cusp QL Ni et al. (2016) 0 = absent, 1 = distolingually positioned relative to protoconid, 2 = lingually positioned 57 P metaconid position QL Ni et al. (2016)* 3 relative to protoconid, 3 = distal to protoconid P talonid cusp 0 = one distinct cusp present, 1 = two distinct cusps present, 2 = polymorphic between two 58 3 QL This Study development and three cusps present, 3 = three distinct cusps present

59 Lower P3 molarization 0 = premolariform, 1 = submolariform, 2 = molariform QL Qi and Beard (1998)

60 P3 shape 0 = short/broad, 1 = long/narrow QN Gilbert et al. (2017)

61 P3 root number 0 = two, 1 = one QL Gilbert et al. (2017)

62 P4 metaconid 0 = absent, 1 = small distinct cusp, 2 = large cusp QL Ni et al. (2016) 0 = slightly distolingually positioned relative to the protoconid, 1 = significatnly distonlingually positioned, 2 = lingually positioned relative to protoconid, 3 = lingually 63 P metaconid position QL Ni et al. (2016) 4 positioned, slightly mesial relative to the protoconid, 4 = mesiodistally inline with the protoconid

P4 protoconid height 64 0 = similar in height, 1 = P4 higher, 2 = P4 significantly higher QN Ni et al. (2016)* relative to that of M1

65 P4 root number 0 = two, 1 = one QL Ni et al. (2016)

66 P1 presence 0 = present, 1 = absent QL Ni et al. (2016)

67 P2 presense 0 = present, 1 = absent QL Ni et al. (2016)

68 P3 evolutionary loss 0 = absent, 1 = present QL Ni et al. (2016)

69 P3 size relative to M1 0 = very small, much smaller than M1, 1 = slightly smaller than M1 QN Ni et al. (2016)* P3 size relative to upper 70 0 = P3 smaller than P4, 1 = P3 similar in size QN Ni et al. (2016)* P4 0 = absent, 1 = present, a small cusp, or a swollen on the postparacrista, 2 = present, large 71 P3 metacone QL Ni et al. (2016) cusp 72 P3 protocone 0= absent, 1 = small cusp, 2 = large cusp QL Ni et al. (2016)

46 73 P3 root number 0 = one, 1 = two, 2 = three QL Ni et al. (2016) P4 size relative to upper 74 0 = much shorter, 1 = shorter, 2 = slightly shorter to similar in size QN Ni et al. (2016)* M1 75 P4 protocone 0 = absent, 1 = present QL Ni et al. (2016)

76 P4 metacone 0 = absent, 1 = small cusp or swollen on the postparacrista, 2 = present as a distinct cusp QL Ni et al. (2016)

77 P4 hypocone 0 = absent, 1 = present QL Ni et al. (2016)

78 P4 root number 0 = three, 1 = two, 2 = one QL Ni et al. (2016) Upper premolar buccal 79 0 = absent/weak, 1 = polymorphic/intermediate, 2 = strong/continuous QL This study cingulum strength Upper premolar lingual 80 0 = absent/weak, 1 = polymorphic/intermediate, 2 = strong/continuous QL This study cingulum strength Lower premolar buccal 81 0 = absent/weak, 1 = polymorphic/intermediate, 2 = strong/continuous QL This study cingulid strength Lower premolar lingual 82 0 = absent/weak, 1 = polymorphic/intermediate, 2 = strong/continuous QL This study cingulid strength *Indicates character was modified from original source

47

a) b)

Figure 9: Summary of “conservative” analysis that examined 22 taxa. Tree length = 216, CI = .409, RI = .342. a) Strict-consensus tree of the 690 MPTs. b) Majority-rule consensus tree of all trees within 50% MPT

48 a) b)

Figure 10: Summary of “liberal” analysis that examined 21 taxa. Tree length = 221, CI = .422, RI = .379. a) Strict-consensus tree of the 31 MPTs. b) Majority-rule consensus tree of all trees within 50% of the MPT

49 a) b)

Figure 11: Summary of “conservative” analysis that examined 19 taxa. Tree length = 196, CI = .475, RI = .409. a) Strict-consensus tree of the 10 MPTs. b) Majority-rule consensus tree of all trees within 50% of the MPT

50 55

Figure 12: Summary of “liberal” analysis that examined 18 taxa. Tree length = 200, CI = .465, RI = .387. One tree recovered, above is the topology recovered for both the strict-consensus tree and majority-rule tree within 50% of the MPT. Bootstrap support values above 50% are provided above a given branch.

51 Table 3: Comparative absolute measurements of the mandible and lower molars for all taxa sampled in analysis.

M1 M1 M1 M1 M1 M2 M2 M2 M3 M3 M3 Taxon Specimen Reference Mand. MD MDTri BLTal Ht. MD MDTri BLTal MD MDTri BLTal Ht. Dashzeveg and GISPS McKenna (1977); Altanius orlovi coll.7, No. 1.2 0.617 1.341 1.09 1.2 - 1.447 1.5 0.618 1.27 - Rose and Krause 20-8 (1984)* Gingerich et al. Altanius orlovi PSS 20-58 1.13 0.678 0.97 1.18 1.17 0.612 1.055 1.62 0.579 0.89 2.167 (1991)* Gingerich et al. Altanius orlovi PSS 20-85 - - - - 1.14 - 1.05 1.58 - 0.92 - (1991) Gingerich et al. Altanius orlovi PSS 20-136 1.13 - 0.92 0.76 1.16 - 1.07 - - - - (1991)* Altanius 1.153 0.648 1.077 1.007 1.168 0.612 1.156 1.567 0.599 1.026 2.167

UCMP Purgatorius unio Clemens (1974)* 2.200 1.049 1.855 1.993 1.050 1.857 1.967 2.413 1.072 1.443 3.716 107406 Purgatorius titusi UM 90207 Buckley (1997)* ------Purgatorius titusi UM 90195 Buckley (1997)* 1.9 0.903 1.4 1.46 ------Purgatorius titusi UM 90184 Buckley (1997)* - - - - 2 0.945 1.4 - - - - Purgatorius titusi UM 90186 Buckley (1997)* ------2.1 0.999 1.1 - UM VP Van Valen and Purgatorius unio - - - - 1.921 0.75 1.551 - - - - 1504 Sloan (1965)* UM VP Van Valen and Purgatorius unio ------2.26 0.743 1.25 - 1506 Sloan (1965)* Purgatorius 2.050 0.976 1.628 1.727 1.657 1.184 1.639 2.259 0.938 1.263 3.716 Szalay (1976); Teilhardina IRSNB Smith et al. 1.798 0.908 1.366 1.38 1.719 0.82 1.47 2.1 0.728 0.94 2.523 belgica M64 (2006)* CM 73229, CM 67856, Teilhardina CM 70435, Beard (2008)* 1.919 0.852 1.28 0.95 1.606 0.818 1.172 2.01 0.808 1.05 - magnoliana CM 70427** Teilhardina IVPP Ni et al. (2004)* 2.12 1.009 1.404 1.07 2.066 0.921 1.404 1.92 0.746 1.01 1.294 asiatica V12358

52 Teilhardina 1.946 0.923 1.350 1.132 1.797 0.853 1.349 2.012 0.761 1.000 1.909 Cantius torresi UM 66143 Gingerich (1995) - - - - 3.43 - - 4.22 1.91 2.4 - Gingerich Cantius torresi UM 83470 3.418 1.634 2.839 2.66 ------6.486 (1986)* UM Gingerich Cantius torresi 3.381 1.563 2.631 2.89 3.35 1.449 2.902 4.1 1.793 2.31 5.891 101958 (1995)* Cantius torresi UM 84367 Gingerich (1995) - - - - 3.2 - 3.06 - - - 6.56 Cantius torresi UM 86132 Gingerich (1995) ------4.22 - 2.44 - Gingerich Cantius torresi UM 87341 3.47 - 2.96 - 3.62 - 3.26 - - - 7.76 (1995)* Cantius torresi UM 87341 Gingerich (1995) - - - - 3.6 - 3.31 4.65 - 2.74 - YPM Rose et al. Cantius ralstoni 4.074 1.907 3.074 3.2 4.122 1.778 3.593 5.06 1.611 2.83 - 23317 (1994)* Rose et al. Cantius ralstoni UW 8842 3.756 2.016 3.142 2.75 3.93 1.827 3.463 - - - - (2011)* Cantius spp. 3.620 1.780 2.929 2.875 3.607 1.685 3.265 4.449 1.771 2.544 6.674 Donrussellia RI 170 Godinot (1981)* 2.4 1.38 1.261 1.96 2.3 1.14 1.614 2.9 0.9 1.42 4.023 provincialis Donrussellia RI 430 Godinot (2015)* 2.352 0.99 1.759 1.86 ------provincialis Donrussellia RI 174 Godinot (1981) 2.4 - 1.9 ------provincialis Donrussellia RI 406 Godinot (1981) 2.3 - 1.8 ------provincialis Donrussellia RI 238 Godinot (1981) - - - - 2.3 - 1.9 - - - - provincialis Donrussellia RI 407 Godinot (1981) - - - - 2.5 - 1.9 - - - - provincialis Donrussellia 2.363 1.185 1.680 1.907 2.367 1.140 1.805 2.900 0.900 1.421 4.023 provincialis Europolemur Bchs 247 Godinot (1988)* 3.93 1.568 2.72 ------klatti Europolemur Bchs 317 Godinot (1988) 4.17 - 2.83 ------klatti Europolemur BUX 80.90 Godinot (1988)* 4.08 1.692 2.58 ------klatti Europolemur BUX 80.91 Godinot (1988) - - - - 4.11 - 2.87 - - - - klatti

53 Europolemur BUX 80.92 Godinot (1988) ------4.72 - 2.86 - klatti Europolemur BUX Godinot (1988) 4.65 - 3.03 - 4.57 - 3.35 5.27 - 2.9 - klatti 80.107 Europolemur BUX 80.99 Godinot (1988) 4.2 - 2.83 ------klatti Europolemur BUX 80.98 Godinot (1988) - - - - 4.39 - 3.16 - - - - klatti Europolemur BUX 80.97 Godinot (1988) ------4.66 - 2.79 - klatti Europolemur BUX Godinot (1988) 4.13 - 2.9 ------klatti 80.103 Europolemur BUX Godinot (1988) 4.24 - 2.82 ------klatti 66.118 Europolemur BUX Godinot (1988) - - - - 4.56 - 3.25 - - - - klatti 80.102 Europolemur BUX Godinot (1988) ------4.79 - 2.88 - klatti 80.105 Europolemur BUX Godinot (1988) ------4.55 - 2.88 - klatti 66.119 Europolemur Bchs 224 Godinot (1988) - - - - 4.42 - 3.43 - - - - klatti Europolemur Bchs 455 Godinot (1988) - - - - 4.31 - 3.24 - - - - klatti Europolemur Bchs 163 Godinot (1988) ------4.79 - 3.02 - klatti Europolemur Bchs 318 Godinot (1988) ------5.1 - 3.13 - klatti Europolemur Bchs 6358 Godinot (1988) ------4.58 - 2.77 - klatti Europolemur Halle Gilbert (2005) 4.214 - 2.643 - 4.143 ------klatti Europolemur 4.202 1.630 2.794 - 4.358 - 3.217 4.808 - 2.904 - klatti Periconodon Bchs 494 Godinot (1988) 2.94 - 2.13 - 2.99 - 2.4 3.75 - 2.32 - huerzeleri Periconodon Bchs 313 Godinot (1988) 3.04 - 2.18 - 3.13 - 2.48 - - - - huerzeleri

54 Periconodon Bchs 270 Godinot (1988) 3.05 - 2.5 ------huerzeleri Periconodon Bchs 495 Godinot (1988)* 2.97 1.181 2.23 - 2.9 1.197 2.31 3 1.089 1.78 - huerzeleri Periconodon Bchs 314 Godinot (1988) 3.18 - 2.41 ------huerzeleri Periconodon 3.036 1.181 2.290 - 3.007 1.197 2.397 3.375 1.089 2.050 - huerzeleri Femenias-Gaul et Agerinia roselli IPS-82793 3.24 1.237 2.553 2.05 ------al. (2016)* Femenias-Gaul et Agerinia roselli IPS-82816 3.24 1.404 2.421 2.15 ------al. (2016)* Femenias-Gaul et Agerinia roselli IPS-2542 - - - - 3.26 1.333 2.474 - - - - al. (2016)* Femenias-Gaul et Agerinia roselli IPS-82794 - - - - 3.51 1.246 2.842 - - - - al. (2016)* Femenias-Gaul et Agerinia roselli IPS-1981 - - - - 3.31 1.176 2.562 3.93 1.098 2.15 - al. (2016)* Femenias-Gaul et Agerinia roselli IPS-2541 - - - - 2.844 1.053 2.614 3.28 1.053 1.91 - al. (2016)* Femenias-Gaul et Agerinia roselli IPS-82795 ------3.97 1.07 2.3 - al. (2016)* Agerinia roselli 3.24 1.321 2.487 2.1 3.231 1.202 2.623 3.73 1.074 2.12 - Protoadapis MNHN Russellet al. - - - - 5.1 - 3.8 5.8 - 3.5 - curvicuspidens AL-5179 (1967) Protoadapis MNHN Russell et al. 4.8 - 3.8 - 5.2 - 3.9 - - - - curvicuspidens AL-5180 (1967) MNHN- Protoadapis Russell et al. Louis-15 4.5 1.828 3.597 3.12 4.7 1.577 3.808 5.6 1.446 3.65 10.414 curvicuspidens (1967)* Ma Protoadapis Russell et al. Lyon 1991 5.5 - 3.9 ------curvicuspidens (1967) Protoadapis Russell et al. AL-5181 - - - - 4.9 - 3.5 - - - - curvicuspidens (1967) Protoadapis Louis-172 Russell et al. 5 - 3.4 ------curvicuspidens Gr (1967) Protoadapis Louis-176 Russell et al. 4.9 - 3.5 ------curvicuspidens Gr (1967)

55 Protoadapis Louis-185 Russell et al. 4.9 - 3.3 ------curvicuspidens Gr (1967) Protoadapis Louis-108 Russell et al. 3.8 - 2.7 ------curvicuspidens Gr (1967) Protoadapis Louis-106 Russell et al. 3.9 - 2.7 ------curvicuspidens Gr (1967) Protoadapis Louis-112 Russell et al. 4.1 - 3.1 ------curvicuspidens Gr (1967) Protoadapis Russell et al. AL-5190 - - - - 5.3 - 3.9 - - - - curvicuspidens (1967) Protoadapis Louis-105 Russell et al. - - - - 4 - 3.2 - - - - curvicuspidens Gr (1967) Protoadapis Louis-99 Russell et al. - - - - 4.1 - 3.2 - - - - curvicuspidens Gr (1967) Protoadapis Louis-112 Russell et al. - - - - 4 - 3.1 - - - - curvicuspidens Gr (1967) Protoadapis Russell et al. AL-5189 ------5.1 - 3.2 - curvicuspidens (1967) Protoadapis Louis-25 Russell et al. ------5.9 - 3.2 - curvicuspidens Mt (1967) Protoadapis Louis-96 Russell et al. ------6.3 - 3.6 - curvicuspidens Gr (1967) Protoadapis Louis-104 Russell et al. ------5 - 2.9 - curvicuspidens Gr (1967) Protoadapis Louis-110 Russell et al. ------5.2 - 2.9 - curvicuspidens Gr (1967) Protoadapis Louis-109 Russell et al. ------4.6 - 2.9 - curvicuspidens Gr (1967) Protoadapis 4.600 1.828 3.333 3.115 4.663 1.577 3.551 5.438 1.446 3.232 10.414 curvicuspidens Marcgodinotius Gu 7 Rose et al. (2009) 2.4 - 1.65 ------indicus Marcgodinotius Rose et al. GU 40 ------indicus (2009)* Marcgodinotius Rose et al. GU 44 2.2 1.008 1.4 1.56 ------indicus (2009)* Marcgodinotius GU 45 Rose et al. (2009) - - - - 2.35 0.932 1.75 - - - - indicus

56 Marcgodinotius GU 46 Rose et al. (2009) ------2.3 0.707 1.35 - indicus Marcgodinotius GU 49 Rose et al. (2009) 2.4 - 1.55 ------indicus Marcgodinotius GU 51 Rose et al. (2009) ------2.25 - 1.3 - indicus Marcgodinotius GU 52 Rose et al. (2009) ------2.4 - 1.3 - indicus Marcgodinotius GU 54 Rose et al. (2009) 2.3 - 1.55 ------indicus Marcgodinotius Rose et al. GU 227 - - - - 2.3 1.025 1.75 2.4 0.981 1.4 - indicus (2009)* Marcgodinotius Rose et al. GU 600 2.3 1 1.55 ------indicus (2009)* Marcgodinotius GU 611 Rose et al. (2009) 2.2 - 1.4 ------indicus Marcgodinotius GU 643 Rose et al. (2009) - - - - 2.15 - 1.55 - - - - indicus Marcgodinotius GU 644 Rose et al. (2009) - - - - 2.2 - 1.5 - - - - indicus Marcgodinotius GU 645 Rose et al. (2009) - - - - 2.4 - 1.7 - - - - indicus Marcgodinotius GU 646 Rose et al. (2009) - - - - 2.15 - 1.55 - - - - indicus Marcgodinotius Rose et al. GU 727 - - - - 2.2 1 1.65 - - - - indicus (2009)* Marcgodinotius Rose et al. GU 743 - - - - 2.2 0.864 1.6 - - - - indicus (2009)* Marcgodinotius Rose et al. GU 744 - - - - 2.3 1.012 1.6 - - - - indicus (2009)* Marcgodinotius GU 1534 Rose et al. (2009) ------2.3 - 1.2 - indicus Marcgodinotius Gu 1575 Rose et al. (2009) - - - - 2.3 - 1.6 - - - - indicus Marcgodinotius Gu 1591 Rose et al. (2009) ------2.35 - 1.3 - indicus Marcgodinotius GU 1602 Rose et al. (2009) - - - - 2.3 - 1.6 - - - - indicus

57 Marcgodinotius IITR 800 Rose et al. (2009) - - - - 2.23 - 1.73 - - - - indicus Marcgodinotius 2.3 1.004 1.517 1.564 2.257 0.967 1.632 2.333 0.844 1.308 - indicus Asiadapis GU/RSR/V Rose et al. 2.85 1.309 2 1.51 2.9 1.082 2.15 - - - 4.3 cambayensis AS-6 (2007)* Asiadapis Rose et al. GU 32 ------2.8 0.925 1.7 - cambayensis (2009)* Asiadapis Rose et al. GU 36 - - - - 2.65 1.222 2.1 - - - - cambayensis (2009)* Asiadapis GU 37 Rose et al. (2009) - - - - 2.85 - 2.35 - - - - cambayensis Asiadapis GU 598 Rose et al. (2009) ------3.4 0.982 2 - cambayensis Asiadapis GU 642 Rose et al. (2009) - - - - 2.95 - 2.3 - - - - cambayensis Asiadapis Rose et al. GU 745 3.1 1.248 2.2 1.57 3.1 1.036 2.4 - - - 4.1 cambayensis (2009)* Asiadapis GU 1505 Rose et al. (2009) ------3.25 - 1.8 - cambayensis Asiadapis GU 1649 Rose et al. (2009) ------3.3 - 1.95 - cambayensis Suratius robustus IITR 928 Rose et al. (2009) - - - - 2.7 - 2.4 - - - - Asiadapis 2.983 1.279 2.067 1.538 2.858 1.113 2.283 3.19 0.954 1.86 4.2 cambayensis Adapoides IVPP Beard et al. - - - - 2.65 1.34 1.845 3.55 1.205 1.68 - troglodytes V11023 (1994)* Adapoides - - - - 2.65 1.34 1.845 3.55 1.205 1.68 - troglodytes GSP-UM Gunnell et al. Panobius russelli - - - - 2.1 0.916 1.5 - - - 3.243 5073 (2008)* GSP-UM Gunnell et al. Panobius russelli 2.1 - 1.5 ------6768 (2008) GSP-UM Gunnell et al. Panobius russelli - - - - 2 - 1.4 - - - - 6769 (2008) GSP-UM Gunnell et al. Panobius russelli 2.2 1.067 1.4 ------6770 (2008)*

58 GSP-UM Gunnell et al. Panobius russelli - - - 2 - 1.4 - - - - 6771 (2008) GSP-UM Gunnell et al. Panobius russelli 2.1 - 1.3 ------6773 (2008) GSP-UM Gunnell et al. Panobius russelli 2.2 - 1.4 ------6774 (2008) GSP-UM Gunnell et al. Panobius russelli - - - - 2 - 1.5 - - - - 6777 (2008) GSP-UM Gunnell et al. Panobius russelli - - - - 2 - 1.5 - - - - 6778 (2008) GSP-UM Gunnell et al. Panobius russelli - - - - 2 - 1.4 - - - - 6779 (2008) GSP-UM Gunnell et al. Panobius russelli ------1.9 - 1.4 - 6780 (2008) GSP-UM Gunnell et al. Panobius russelli ------2.1 - 1.4 - 6781 (2008) GSP-UM Gunnell et al. Panobius russelli ------2.2 - 1.4 - 6782 (2008) Panobius russelli 2.15 1.067 1.4 - 2.017 0.916 1.45 2.067 1.4 3.243 Gilbert et al. Hoanghonius Unnumbere (2017); Gingerich - - - - 4 1.5 3.3 4.6 1.3 2.7 7.9 stehlini d, Uppsala et al (1994) Hoanghonius IVPP Tong et al. 3.9 1.989 2.8 2.03 4 3 4.6 2.8 7.2 stehlini 10220 (1999)* Hoanghonius 3.9 1.989 2.8 2.03 4 1.5 3.15 4.6 1.3 2.75 7.55 stehlini Gilbert et al. Rencunius zhoui IVPP 5312 (2017); Gingerich 3.8 1.9 3 2.71 4.2 1.7 3.5 - - - - et al (1994) Gingerich et al. Rencunius zhoui IVPP 5311 - - - - 4 - 3.9 - - - - (1994) Specimen Woo and Chow Rencunius zhoui ------4.5 - 2.9 - No. 3 (1957) Rencunius zhoui 3.8 1.9 3 2.71 4.1 1.7 3.7 4.5 - 2.9 - Gilbert et al. Wailekia TF 2632 (2017); Ducroq et - - - - 4.6 1.6 3.6 4.7 1.5 3.2 - orientale al. (1995)*

59 Wailekia - - - - 4.6 1.6 3.6 4.7 1.5 3.2 - orientale Paukkaungia Beard et al. NMMP 55 3.59 1.369 2.45 2.3 ------parva (2007)* Paukkaungia Beard et al. NMMP 57 - - - - 3.31 1.293 2.41 - - - - parva (2007)* Paukkaungia 3.59 1.369 2.45 2.3 3.31 1.293 2.41 - - - - parva Kyitchaungia Beard et al. NMMP 28 - - - - 4.39 - 3 - - - - takaii (2007)* Kyitchaungia - - - - 4.39 - 3 - - - - takaii Guangxilemur Marivaux et al. DBC 2170 5.067 - 3.506 ------singsilai (2002) Guangxilemur Marivaux et al. DBC 2171 5.028 - 3.606 ------singsilai (2002) Guangxilemur 5.048 - 3.556 ------singsilai MD = maximum mesiodistal length, MDTri = maximum mesiodistal length of trigonid, BLTal = maximum buccal-lingual width of talonid, Ht. = crown height at protoconid, *Indicates additional novel measurements were recorded from published photos using ImageJ method described above, **Indicates composite specimens, ***Indicates a species was synonymized with another genus/species. Population averages in bold. All measurements in millimeters.

60 Table 4: Comparative shape and area values of lower molars derived from absolute lower molar measurements

M3 M1 M1 M1 M2 M2 M2 M3 M3 Taxon Specimen Reference shapeTr shape shapeTri area shape shapeTri area shape area i Dashzeveg and GISPS coll.7, McKenna (1977); Altanius orlovi 0.895 0.514 1.609 0.829 - 1.736 1.186 0.412 1.898 No. 20-8 Rose and Krause (1984)* Gingerich et al. Altanius orlovi PSS 20-58 1.165 0.600 1.096 1.109 0.523 1.234 1.812 0.357 1.448 (1991)* Gingerich et al. Altanius orlovi PSS 20-85 - - - 1.086 - 1.197 1.717 - 1.454 (1991) Gingerich et al. Altanius orlovi PSS 20-136 1.228 - 1.040 1.084 - 1.241 - - - (1991)* Altanius 1.096 0.557 1.248 1.027 0.523 1.352 1.572 0.385 1.600 UCMP Purgatorius unio Clemens (1974)* 1.186 0.477 4.081 0.534 1.769 2.065 1.672 0.444 3.482 107406 Purgatorius titusi UM 90195 Buckley (1997)* 1.357 0.475 2.660 ------Purgatorius titusi UM 90184 Buckley (1997)* - - - 1.429 0.473 2.800 - - - Purgatorius titusi UM 90186 Buckley (1997)* ------1.909 0.476 2.310 Van Valen and Purgatorius unio UM VP 1504 - - - 1.239 0.390 2.979 - - - Sloan (1965)* Van Valen and Purgatorius unio UM VP 1506 ------1.818 0.328 2.819 Sloan (1965)* Purgatorius 1.272 0.476 3.371 1.067 0.877 2.615 1.800 0.416 2.870 Szalay (1976); Teilhardina belgica IRSNB M64 Smith et al. 1.316 0.505 2.456 1.169 0.477 2.527 2.232 0.347 1.976 (2006)* CM 73229, CM 67856, Teilhardina magnoliana Beard (2008)* 1.499 0.444 2.456 1.370 0.509 1.882 1.916 0.401 2.117 CM 70435, CM 70427** Teilhardina asiatica IVPP V12358 Ni et al. (2004)* 1.510 0.476 2.976 1.472 0.446 2.901 1.904 0.388 1.938 Teilhardina 1.442 0.475 2.630 1.337 0.477 2.437 2.017 0.379 2.010 Cantius torresi UM 66143 Gingerich (1995) ------1.758 0.453 10.128 Cantius torresi UM 83470 Gingerich (1986)* 1.204 0.478 9.704 ------Cantius torresi UM 101958 Gingerich (1995)* 1.285 0.462 8.895 1.154 0.433 9.722 1.777 0.437 9.459

61 Cantius torresi UM 84367 Gingerich (1995) - - - 1.046 - 9.792 - - - Cantius torresi UM 86132 Gingerich (1995) ------1.730 - 10.297 11.80 Cantius torresi UM 87341 Gingerich (1995)* 1.172 - 10.271 1.110 - - - - 1 11.91 Cantius torresi UM 87341 Gingerich (1995) - - - 1.088 - 1.697 - 12.741 6 14.81 Cantius ralstoni YPM 23317 Rose et al. (1994)* 1.325 0.468 12.523 1.147 0.431 1.785 0.319 14.324 0 13.61 Cantius ralstoni UW 8842 Rose et al. (2011)* 1.195 0.537 11.801 1.135 0.465 - - - 0 11.94 Cantius spp. 1.236 0.486 10.639 1.113 0.443 1.749 0.403 11.390 2 Donrussellia provincialis RI 170 Godinot (1981)* 1.903 0.575 3.026 1.425 0.496 3.712 2.041 0.310 4.121 Donrussellia provincialis RI 430 Godinot (2015)* 1.337 0.421 4.137 ------Donrussellia provincialis RI 174 Godinot (1981) 1.263 - 4.560 ------Donrussellia provincialis RI 406 Godinot (1981) 1.278 - 4.140 ------Donrussellia provincialis RI 238 Godinot (1981) - - - 1.211 - 4.370 - - - Donrussellia provincialis RI 407 Godinot (1981) - - - 1.316 - 4.750 - - - Donrussellia provincialis 1.445 0.498 3.966 1.317 0.496 4.277 2.041 0.310 4.121 Europolemur klatti Bchs 247 Godinot (1988)* 1.445 0.399 10.690 ------Europolemur klatti Bchs 317 Godinot (1988) 1.473 - 11.801 ------Europolemur klatti BUX 80.90 Godinot (1988)* 1.581 0.415 10.526 ------11.79 Europolemur klatti BUX 80.91 Godinot (1988) - - - 1.432 - - - - 6 Europolemur klatti BUX 80.92 Godinot (1988) ------1.650 - 13.499 15.31 Europolemur klatti BUX 80.107 Godinot (1988) 1.535 - 14.090 1.364 - 1.817 - 15.283 0 Europolemur klatti BUX 80.99 Godinot (1988) 1.484 - 11.886 ------13.87 Europolemur klatti BUX 80.98 Godinot (1988) - - - 1.389 - - - - 2 Europolemur klatti BUX 80.97 Godinot (1988) ------1.670 - 13.001 Europolemur klatti BUX 80.103 Godinot (1988) 1.424 - 11.977 ------Europolemur klatti BUX 66.118 Godinot (1988) 1.504 - 11.957 ------Europolemur klatti BUX 80.102 Godinot (1988) - - - 1.403 - 14.82 - - - Europolemur klatti BUX 80.105 Godinot (1988) ------1.663 - 13.795 Europolemur klatti BUX 66.119 Godinot (1988) ------1.580 - 13.104

62 15.16 Europolemur klatti Bchs 224 Godinot (1988) - - - 1.289 - - - - 1 13.96 Europolemur klatti Bchs 455 Godinot (1988) - - - 1.330 - - - - 4 Europolemur klatti Bchs 163 Godinot (1988) ------1.586 - 14.466 Europolemur klatti Bchs 318 Godinot (1988) ------1.629 - 15.963 Europolemur klatti Bchs 6358 Godinot (1988) ------1.653 - 12.687 Europolemur klatti Halle Gilbert (2005) 1.595 - 11.138 ------14.15 Europolemur klatti 1.505 0.407 11.758 1.368 - 1.656 - 13.975 4 Periconodon huerzeleri Bchs 494 Godinot (1988) 1.380 - 6.262 1.246 - 7.176 1.616 - 8.700 Periconodon huerzeleri Bchs 313 Godinot (1988) 1.394 - 6.627 1.262 - 7.762 - - - Periconodon huerzeleri Bchs 270 Godinot (1988) 1.220 - 7.625 ------Periconodon huerzeleri Bchs 495 Godinot (1988)* 1.332 0.398 6.623 1.255 0.413 6.699 1.685 0.363 5.340 Periconodon huerzeleri Bchs 314 Godinot (1988) 1.320 - 7.664 ------Periconodon huerzeleri 1.329 0.398 6.960 1.254 0.413 7.212 1.651 0.363 7.020 Femenias-Gaul et Agerinia roselli IPS-82793 1.269 0.382 8.272 ------al. (2016)* Femenias-Gaul et Agerinia roselli IPS-82816 1.338 0.433 7.844 ------al. (2016)* Femenias-Gaul et Agerinia roselli IPS-2542 - - - 1.318 0.409 8.065 - - - al. (2016)* Femenias-Gaul et Agerinia roselli IPS-82794 - - - 1.235 0.355 9.975 - - - al. (2016)* Femenias-Gaul et Agerinia roselli IPS-1981 - - - 1.292 0.355 8.480 1.829 0.279 8.446 al. (2016)* Femenias-Gaul et Agerinia roselli IPS-2541 - - - 1.088 0.370 7.434 1.715 0.321 6.277 al. (2016)* Femenias-Gaul et Agerinia roselli IPS-82795 ------1.728 0.270 9.123 al. (2016)* Agerinia roselli 1.304 0.408 8.058 1.233 0.372 8.489 1.757 0.290 7.948 Protoadapis MNHN AL- Russell et al. 19.38 - - - 1.342 - 1.657 - 20.300 curvicuspidens 5179 (1967) 0 Protoadapis MNHN AL- Russell et al. 20.28 1.263 18.240 1.333 - - - - curvicuspidens 5180 (1967) 0 Protoadapis MNHN-Louis- Russell et al. 17.89 1.251 0.406 16.187 1.234 0.336 1.533 0.258 20.462 curvicuspidens 15 Ma (1967)* 8

63 Protoadapis Russell et al. Lyon 1991 1.410 - 21.450 ------curvicuspidens (1967) Protoadapis Russell et al. 17.15 AL-5181 - - - 1.400 - - - - curvicuspidens (1967) 0 Protoadapis Russell et al. Louis-172 Gr 1.471 - 17.000 ------curvicuspidens (1967) Protoadapis Russell et al. Louis-176 Gr 1.400 - 17.150 ------curvicuspidens (1967) Protoadapis Russell et al. Louis-185 Gr 1.485 - 16.170 ------curvicuspidens (1967) Protoadapis Russell et al. Louis-108 Gr 1.407 - 10.260 ------curvicuspidens (1967) Protoadapis Russell et al. Louis-106 Gr 1.444 - 10.530 ------curvicuspidens (1967) Protoadapis Russell et al. Louis-112 Gr 1.323 - 12.710 ------curvicuspidens (1967) Protoadapis Russell et al. 20.67 AL-5190 - - - 1.359 - - - - curvicuspidens (1967) 0 Protoadapis Russell et al. 12.80 Louis-105 Gr - - - 1.250 - - - - curvicuspidens (1967) 0 Protoadapis Russell et al. 13.12 Louis-99 Gr - - - 1.281 - - - - curvicuspidens (1967) 0 Protoadapis Russell et al. 12.40 Louis-112 Gr - - - 1.290 - - - - curvicuspidens (1967) 0 Protoadapis Russell et al. AL-5189 ------1.594 - 16.32 curvicuspidens (1967) Protoadapis Russell et al. Louis-25 Mt ------1.844 - 18.88 curvicuspidens (1967) Protoadapis Russell et al. Louis-96 Gr ------1.75 - 22.68 curvicuspidens (1967) Protoadapis Russell et al. Louis-104 Gr ------1.724 - 14.5 curvicuspidens (1967) Protoadapis Russell et al. Louis-110 Gr ------1.793 - 15.08 curvicuspidens (1967) Protoadapis Russell et al. Louis-109 Gr ------1.586 - 13.34 curvicuspidens (1967)

64 Protoadapis 16.71 1.384 0.406 15.522 1.311 0.336 1.685 0.258 17.695 curvicuspidens 2 Marcgodinotius indicus Gu 7 Rose et al. (2009) 1.455 - 3.960 ------Marcgodinotius indicus GU 44 Rose et al. (2009)* 1.571 0.458 3.080 ------Marcgodinotius indicus GU 45 Rose et al. (2009) - - - 1.343 0.397 4.113 - - - Marcgodinotius indicus GU 46 Rose et al. (2009) ------1.704 0.307 3.105 Marcgodinotius indicus GU 49 Rose et al. (2009) 1.548 - 3.720 ------Marcgodinotius indicus GU 51 Rose et al. (2009) ------1.731 - 2.925 Marcgodinotius indicus GU 52 Rose et al. (2009) ------1.846 - 3.120 Marcgodinotius indicus GU 54 Rose et al. (2009) 1.484 - 3.565 ------Marcgodinotius indicus GU 227 Rose et al. (2009)* - - - 1.314 0.446 4.025 1.714 0.409 3.360 Marcgodinotius indicus GU 600 Rose et al. (2009)* 1.484 0.435 3.565 ------Marcgodinotius indicus GU 611 Rose et al. (2009) 1.571 - 3.080 ------Marcgodinotius indicus GU 643 Rose et al. (2009) - - - 1.387 - 3.333 - - - Marcgodinotius indicus GU 644 Rose et al. (2009) - - - 1.467 - 3.300 - - - Marcgodinotius indicus GU 645 Rose et al. (2009) - - - 1.412 - 4.080 - - - Marcgodinotius indicus GU 646 Rose et al. (2009) - - - 1.387 - 3.333 - - - Marcgodinotius indicus GU 703 Rose et al. (2009) ------Marcgodinotius indicus GU 727 Rose et al. (2009)* - - - 1.333 0.455 3.630 - - - Marcgodinotius indicus GU 743 Rose et al. (2009)* - - - 1.375 0.393 3.520 - - - Marcgodinotius indicus GU 744 Rose et al. (2009)* - - - 1.438 0.440 3.680 - - - Marcgodinotius indicus GU 1534 Rose et al. (2009) ------1.917 - 2.760 Marcgodinotius indicus Gu 1575 Rose et al. (2009) - - - 1.438 - 3.680 - - - Marcgodinotius indicus Gu 1591 Rose et al. (2009) ------1.808 - 3.055 Marcgodinotius indicus GU 1602 Rose et al. (2009) - - - 1.438 - 3.680 - - - Marcgodinotius indicus IITR 800 Rose et al. (2009) - - - 1.289 - 3.858 - - - Marcgodinotius indicus 1.519 0.446 3.495 1.385 0.426 3.686 1.787 0.358 3.054 GU/RSR/VAS Asiadapis cambayensis Rose et al. (2007)* 1.425 0.459 5.700 1.349 0.373 6.235 - - - -6 Asiadapis cambayensis GU 32 Rose et al. (2009)* ------1.647 0.330 4.760 Asiadapis cambayensis GU 35 Rose et al. (2009)* ------Asiadapis cambayensis GU 36 Rose et al. (2009)* - - - 1.262 0.461 5.565 - - - Asiadapis cambayensis GU 37 Rose et al. (2009) - - - 1.213 - 6.698 - - - Asiadapis cambayensis GU 598 Rose et al. (2009) ------1.700 0.289 6.800 Asiadapis cambayensis GU 642 Rose et al. (2009) - - - 1.283 - 6.785 - - - Asiadapis cambayensis GU 745 Rose et al. (2009)* 1.409 0.403 6.820 1.292 0.334 7.440 - - -

65 Asiadapis cambayensis GU 1505 Rose et al. (2009) ------1.806 - 5.850 Asiadapis cambayensis GU 1537 Rose et al. (2009) 1.500 - 6.000 ------Asiadapis cambayensis GU 1649 Rose et al. (2009) ------1.692 - 6.435 Suratius robustus IITR 928 Rose et al. (2009) - - - 1.125 - 6.480 - - - Asiadapis cambayensis 1.445 0.431 6.173 1.254 0.389 6.534 1.711 0.310 5.961 Beard et al. Adapoides troglodytes IVPP V11023 - - - 1.436 0.506 4.889 2.112 0.339 5.968 (1994)* Adapoides troglodytes - - - 1.436 0.506 4.889 2.112 0.339 5.968 Gunnell et al. Panobius russelli GSP-UM 5073 - - - 1.400 0.436 3.150 - - - (2008)* Gunnell et al. Panobius russelli GSP-UM 6768 1.400 - 3.150 ------(2008) Gunnell et al. Panobius russelli GSP-UM 6769 - - - 1.429 - 2.8 - - - (2008) Gunnell et al. Panobius russelli GSP-UM 6770 1.571 0.485 3.080 - - - - - (2008)* Gunnell et al. Panobius russelli GSP-UM 6771 - - - 1.429 - 2.8 - - - (2008) Gunnell et al. Panobius russelli GSP-UM 6773 1.615 - 2.730 ------(2008) Gunnell et al. Panobius russelli GSP-UM 6774 1.571 - 3.080 ------(2008) Gunnell et al. Panobius russelli GSP-UM 6777 - - - 1.333 - 3 - - - (2008) Gunnell et al. Panobius russelli GSP-UM 6778 - - - 1.333 - 3 - - - (2008) Gunnell et al. Panobius russelli GSP-UM 6779 - - - 1.429 - 2.8 - - - (2008) Gunnell et al. Panobius russelli GSP-UM 6780 ------1.357 - 2.66 (2008) Gunnell et al. Panobius russelli GSP-UM 6781 ------1.5 - 2.94 (2008) Gunnell et al. Panobius russelli GSP-UM 6782 ------1.571 - 3.08 (2008) Panobius russelli 1.540 0.485 3.010 1.392 0.436 2.925 1.476 - 2.893

66 Gilbert et al. Unnumbered, 13.20 Hoanghonius stehlini (2017); Gingerich - - - 1.212 0.375 1.704 0.283 12.420 Uppsala 0 et al (1994) Hoanghonius stehlini IVPP 10220 Tong et al. (1999)* 1.393 0.510 10.920 1.333 - 12.00 1.643 - 12.880 Hoanghonius stehlini 1.393 0.510 10.920 1.273 0.375 12.60 1.673 0.283 12.650 Gilbert et al. 14.70 Rencunius zhoui IVPP 5312 (2017); Gingerich 1.267 0.500 11.400 1.200 0.405 - - - 0 et al (1994) Gingerich et al. Rencunius zhoui IVPP 5311 - - - 1.026 - 15.60 - - - (1994) Specimen No. Woo and Chow Rencunius zhoui ------1.552 - 13.050 3 (1957) Rencunius zhoui 1.267 0.500 11.400 1.113 0.405 15.15 1.552 0.291 13.050 Gilbert et al. Wailekia orientale TF 2632 (2017); Ducroq et - - - 1.278 0.348 16.56 1.469 0.319 15.040 al. (1995)* Wailekia orientale - - - 1.278 0.348 16.56 1.469 0.319 15.040 Beard et al. Paukkaungia parva NMMP 55 1.465 0.381 8.796 ------(2007)* Beard et al. Paukkaungia parva NMMP 57 - - - 1.373 0.391 7.977 - - - (2007)* Paukkaungia parva 1.465 0.381 8.796 1.373 0.391 7.977 - - - Beard et al. Kyitchaungia takaii NMMP 28 - - - 1.463 0.410 13.17 - - - (2007)* Kyitchaungia takaii - - - 1.463 0.410 13.17 - - - Marivaux et al. Guangxilemur singsilai DBC 2170 1.445 ------(2002) Marivaux et al. Guangxilemur singsilai DBC 2171 1.394 ------(2002) Guangxilemur singsilai 1.420 Shape = MD/BLTal, Area = MD*BLTal, ShapeTri = MDTri/MD (i.e., ShapeTri measures degree of trigonid compression), *Indicates additional novel measurements were recorded from published photos using ImageJ method described above, **Indicates composite specimens, ***Indicates a species was synonymized with another genus/species. Population averages in bold. All measurements in millimeters.

67 Table 5: Comparative absolute measurements of the lower premolars for all taxa sampled in analysis

P2 P3 P4 P4 Taxon Specimen Reference P1 MD P1 BL P2 MD P2 BL P3 MD P3 Ht. P4 MD Ht. BL BL Ht.

Dashzeveg and GISPS coll.7, McKenna (1977); Altanius orlovi ------1 1.067 1.162 No. 20-8 Rose and Krause (1984)* Gingerich et al. Altanius orlovi PSS 20-58 - - - - - 0.96 0.72 1.235 0.92 0.87 1.261 (1991)* Gingerich et al. Altanius orlovi PSS 20-136 - - - - - 0.78 0.71 0.86 0.94 0.8 0.84 (1991)* Altanius - - - - - 0.870 0.715 1.048 0.953 0.912 1.088 UCMP Purgatorius unio Clemens (1974)* - - 1.228 0.736 1.515 1.387 0.924 1.667 1.832 1.233 2.406 107406 Purgatorius titusi UM 90207 Buckley (1997)* ------1.6 1.1 1.408 Van Valen and Purgatorius unio UM VP 1616 ------1.636 1.113 1.86 Sloan (1965)* Purgatorius - - 1.228 0.736 1.515 1.387 0.924 1.667 1.689 1.149 1.891 Szalay (1976); Teilhardina IRSNB M64 Smith et al. - - - - - 1.328 0.936 1.473 1.355 1.106 1.744 belgica (2006)* CM 73229, Teilhardina CM 67856, Beard (2008)* ------1.238 1.172 - magnoliana CM 70435, CM 70427** Teilhardina IVPP V12358 Ni et al. (2004)* 0.6 0.5 0.857 0.592 0.77 1.565 1.009 1.454 1.608 1.303 1.773 asiatica Teilhardina 0.585 0.482 0.857 0.592 0.771 1.447 0.973 1.464 1.400 1.194 1.759 Cantius torresi UM 83470 Gingerich (1986)* - - - - - 2.42 1.8 3.017 2.882 2.379 - Cantius torresi UM 101958 Gingerich (1995)* 1.1 1 1.488 1.131 2.21 2.269 1.665 2.733 2.961 2.116 3.191 Cantius torresi UM 87341 Gingerich (1995)* - 1.2 1.82 1.29 - 2.51 1.84 2.94 2.92 2.48 3.25 Rose et al. Cantius ralstoni YPM 23317 - - - - - 3.196 2.056 4 3.752 2.667 3.89 (1994)* Rose et al. Cantius ralstoni UW 8842 - - - - - 3.112 1.885 2.911 3.492 2.396 3.446 (2011)* Cantius spp. 1.050 1.064 1.654 1.211 2.212 2.701 1.849 3.120 3.201 2.408 3.444

68 Donrussellia RI 430 Godinot (2015)* - - - - - 1.715 1.11 2.331 1.98 1.418 2.364 provincialis Donrussellia RI 396 Godinot (1981) - - - - - 1.9 1 - - - - provincialis Donrussellia RI 175 Godinot (1981) ------2.2 1.5 - provincialis Donrussellia provincialis - - - - - 1.808 1.055 2.331 2.090 1.459 2.364 Europolemur Bchs 247 Godinot (1988)* - - 1.98 1.29 - 3.28 1.89 - - - - klatti Europolemur BUX 80.89 Godinot (1988) ------4.07 2.53 - klatti Europolemur BUX 80.94 Godinot (1988) ------4.2 2.67 - klatti Europolemur BUX 80.100 Godinot (1988) ------3.89 2.56 - klatti Europolemur BUX 80.101 Godinot (1988) ------3.93 2.52 - klatti Europolemur BUX 80.93 Godinot (1988) - - - - - 3.86 2.19 - - - - klatti Europolemur - - 1.980 1.290 - 3.570 2.040 - 4.023 2.570 - klatti Periconodon Bchs 495 Godinot (1988)* ------2.67 1.54 - huerzeleri Periconodon ------2.670 1.540 - huerzeleri Femenias-Gaul et Agerinia roselli IPS-2543 - - - - - 2.27 1.43 2.65 2.77 1.7 2.66 al. (2016)* Agerinia roselli - - - - - 2.27 1.43 2.65 2.77 1.7 2.66 Protoadapis MNHN AL- Russell et al. - - 3.1 2 - 5.2 3.3 - 5.5 3.6 - curvicuspidens 5180 (1967) Protoadapis MNHN- Russell et al. - - - - - 4.3 2.8 5.011 4.9 3.3 4.774 curvicuspidens Louis-15 Ma (1967)* Protoadapis Russell et al. Louis-197 Gr - - - - - 4.7 3 - - - - curvicuspidens (1967) Protoadapis Russell et al. Louis-196 Gr ------4.7 3.1 - curvicuspidens (1967)

69 Protoadapis Russell et al. Lyon 1991 ------5.3 3.6 - curvicuspidens (1967) Protoadapis Russell et al. AL-5181 ------4.4 2.9 - curvicuspidens (1967) Protoadapis curvicuspidens - - 3.1 2 - 4.733 3.033 5.011 4.960 3.300 4.774 Marcgodinotius Rose et al. GU 40 ------2.25 1.3 2.158 indicus (2009)* Marcgodinotius Rose et al. GU 227 - - - - - 1.9 1.25 - 2.3 1.4 - indicus (2009)* Marcgodinotius GU 703 Rose et al. (2009) - - - - - 1.9 1.15 2.339 - - - indicus Marcgodinotius Rose et al. GU 727 ------2 1.35 2.046 indicus (2009)* Marcgodinotius GU 1536 Rose et al. (2009) ------2.3 1.25 - indicus Marcgodinotius GU 1538 Rose et al. (2009) - - - - - 2 1.1 - - - - indicus Marcgodinotius Gu 1544 Rose et al. (2009) - - - - - 2 1.25 - - - - indicus Marcgodinotius GU 1554 Rose et al. (2009) ------2.2 1.3 - indicus Marcgodinotius - - - - - 1.95 1.187 2.339 2.21 1.32 2.102 indicus Asiadapis GU/RSR/VA Rose et al. - - - - - 2.6 1.7 2.321 2.6 1.7 2.114 cambayensis S-6 (2007)* Asiadapis Rose et al. GU 35 - - - - - 2.7 1.5 2.931 - - - cambayensis (2009)* Asiadapis GU 38 Rose et al. (2009) ------2.9 1.7 - cambayensis Asiadapis Rose et al. GU 745 - - - - - 2.95 1.9 2.614 3 1.8 2.404 cambayensis (2009)* Asiadapis GU 1627 Rose et al. (2009) ------2.7 1.65 - cambayensis Suratius robustus IITR 928 Rose et al. (2009) - - - - - 2.68 1.54 - 2.54 1.92 - Asiadapis - - - - - 2.733 1.66 2.622 2.748 1.754 2.259 cambayensis

70 GSP-UM Gunnell et al. Panobius russelli ------1.9 1.3 1.608 5073 (2008)* GSP-UM Gunnell et al. Panobius russelli ------2.2 1.3 - 6775 (2008) Panobius russelli ------2.05 1.3 1.608 Hoanghonius Tong et al. IVPP 10220 - - 2.1 1.4 2.55 3 1.9 3.24 3.4 2.3 2.94 stehlini (1999)* Hoanghonius - - 2.1 1.4 2.55 3 1.9 3.24 3.4 2.3 2.94 stehlini Gilbert et al. Rencunius zhoui IVPP 5312 (2017); Gingerich ------3.6 2.5 3.05 et al (1994) Rencunius zhoui ------3.6 2.5 3.05 Paukkaungia Beard et al. NMMP 56 ------3.54 2.2 2.06 parva (2007)* Paukkaungia Beard et al. NMMP 54 - - - - - 2.53 1.52 1.559 - - - parva (2007)* Paukkaungia - - - - - 2.53 1.52 1.559 3.54 2.2 2.06 parva MD = maximum mesiodistal length, BL = maximum buccal-lingual width of talonid, Ht. = crown height at protoconid, *Indicates additional novel measurements were recorded from published photos using ImageJ method described above, **Indicates composite specimens, ***Indicates a species was synonymized with another genus/species. Population averages in bold. All measurements in millimeters.

71 Table 6: Comparative shape and area values of lower premolars derived from absolute lower molar measurements.

P2 P3 Taxon Specimen Reference P1 area P2 shape P3 area P4 area P4 shape area shape Dashzeveg and GISPS coll.7, No. McKenna (1977); Altanius orlovi - - - - - 1.067 0.937 20-8 Rose and Krause (1984)* Gingerich et al. Altanius orlovi PSS 20-58 - - - 0.691 1.333 0.800 1.057 (1991)* Altanius orlovi PSS 20-85 Gingerich et al. (1991) ------Gingerich et al. Altanius orlovi PSS 20-136 - - - 0.554 1.099 0.752 1.175 (1991)* Altanius - - - 0.623 1.216 0.873 1.057 Purgatorius unio UCMP 107406 Clemens (1974)* - 0.904 1.668 1.282 1.501 2.259 1.486 Purgatorius titusi UM 90207 Buckley (1997)* - - - - - 1.760 1.455 Van Valen and Sloan Purgatorius unio UM VP 1616 - - - - - 1.821 1.470 (1965)* Purgatorius - 0.904 1.668 1.282 1.501 1.947 1.470 Szalay (1976); Smith Teilhardina belgica IRSNB M64 - - - 1.243 1.419 1.499 1.225 et al. (2006)* CM 73229, CM Teilhardina magnoliana 67856, CM 70435, Beard (2008)* - - - - - 1.451 1.056 CM 70427** Teilhardina asiatica IVPP V12358 Ni et al. (2004)* 0.282 0.507 1.448 1.579 1.551 2.095 1.234 Teilhardina 0.282 0.507 1.448 1.411 1.485 1.682 1.172 Cantius torresi UM 83470 Gingerich (1986)* - - - 4.356 1.344 6.856 1.211 Cantius torresi UM 101958 Gingerich (1995)* 1.026 1.683 1.316 3.778 1.363 6.265 1.399 Cantius torresi UM 87341 Gingerich (1995)* - 2.348 1.411 4.618 1.364 7.242 1.177 Cantius ralstoni YPM 23317 Rose et al. (1994)* - - - 6.571 1.554 10.007 1.407 Cantius ralstoni UW 8842 Rose et al. (2011)* - - - 5.866 1.651 8.367 1.457 Cantius spp. 1.026 2.015 1.363 5.038 1.455 7.747 1.330 Donrussellia provincialis RI 430 Godinot (2015)* - - - 1.904 1.545 2.808 1.396 Donrussellia provincialis RI 396 Godinot (1981) - - - 1.900 1.900 - - Donrussellia provincialis RI 175 Godinot (1981) - - - - - 3.300 1.467 Donrussellia provincialis - - - 1.902 1.723 3.054 1.431

72 Europolemur klatti Bchs 247 Godinot (1988)* - 2.554 1.535 6.199 1.735 - - Europolemur klatti BUX 80.89 Godinot (1988) - - - - - 10.297 1.609 Europolemur klatti BUX 80.94 Godinot (1988) - - - - - 11.214 1.573 Europolemur klatti BUX 80.100 Godinot (1988) - - - - - 9.958 1.520 Europolemur klatti BUX 80.101 Godinot (1988) - - - - - 9.904 1.560 Europolemur klatti BUX 80.93 Godinot (1988) - - - 8.453 1.763 - - Europolemur klatti - 2.554 1.535 7.326 1.749 10.343 1.565 Periconodon huerzeleri Bchs 495 Godinot (1988)* - - - - - 4.112 1.734 Periconodon huerzeleri - - - - - 4.112 1.734 Femenias-Gaul et al. Agerinia roselli IPS-2543 - - - 3.246 1.587 4.709 1.629 (2016)* Agerinia roselli - - - 3.246 1.587 4.709 1.629 Protoadapis curvicuspidens MNHN AL-5180 Russell et al. (1967) - 6.2 1.55 17.160 1.576 19.800 1.528 MNHN-Louis-15 Protoadapis curvicuspidens Russell et al. (1967)* - - - 12.040 1.536 16.170 1.485 Ma Protoadapis curvicuspidens Louis-197 Gr Russell et al. (1967) - - - 14.100 1.567 - -

Protoadapis curvicuspidens Louis-196 Gr Russell et al. (1967) - - - - - 14.570 1.516

Protoadapis curvicuspidens Lyon 1991 Russell et al. (1967) - - - - - 19.080 1.472

Protoadapis curvicuspidens AL-5181 Russell et al. (1967) - - - - - 12.760 1.517 Protoadapis curvicuspidens - 6.200 1.550 14.433 1.559 16.476 1.504 Marcgodinotius indicus GU 40 Rose et al. (2009)* - - - - - 2.925 1.731 Marcgodinotius indicus GU 227 Rose et al. (2009)* - - - 2.375 1.520 3.220 1.643 Marcgodinotius indicus GU 703 Rose et al. (2009) - - - 2.185 1.652 - - Marcgodinotius indicus GU 727 Rose et al. (2009)* - - - - - 2.700 1.481 Marcgodinotius indicus GU 1536 Rose et al. (2009) - - - - - 2.875 1.840 Marcgodinotius indicus GU 1538 Rose et al. (2009) - - - 2.200 1.818 - - Marcgodinotius indicus Gu 1544 Rose et al. (2009) - - - 2.500 1.600 - - Marcgodinotius indicus GU 1554 Rose et al. (2009) - - - - - 2.860 1.692 Marcgodinotius indicus - - - 2.315 1.648 2.916 1.677 Asiadapis cambayensis GU/RSR/VAS-6 Rose et al. (2007)* - - - 4.420 1.529 4.420 1.529 Asiadapis cambayensis GU 35 Rose et al. (2009)* - - - 4.050 1.800 - -

73 Asiadapis cambayensis GU 38 Rose et al. (2009) - - - - - 4.930 1.706 Asiadapis cambayensis GU 745 Rose et al. (2009)* - - - 5.605 1.553 5.400 1.667 Asiadapis cambayensis GU 1627 Rose et al. (2009) - - - - - 4.455 1.636 Suratius robustus IITR 928 Rose et al. (2009) - - - 4.127 1.740 4.877 1.323 Asiadapis cambayensis - - - 4.551 1.656 4.816 1.572 Panobius russelli GSP-UM 5073 Gunnell et al. (2008)* - - - - - 2.470 1.462

Panobius russelli GSP-UM 6775 Gunnell et al. (2008) - - - - - 2.860 1.692 Panobius russelli - - - - - 2.665 1.577 Unnumbered, Gilbert et al. (2017); Hoanghonius stehlini ------Uppsala Gingerich et al (1994) Hoanghonius stehlini IVPP 10220 Tong et al. (1999)* 2.940 1.500 5.700 1.579 7.820 1.478 Hoanghonius stehlini - 2.940 1.500 5.700 1.579 7.820 1.478 Gilbert et al. (2017); Rencunius zhoui IVPP 5312 9.000 1.440 Gingerich et al (1994) Rencunius zhoui - - - - - 9.000 1.440 Paukkaungia parva NMMP 56 Beard et al. (2007)* - - - - - 7.788 1.609 Paukkaungia parva NMMP 55 Beard et al. (2007)* ------Paukkaungia parva NMMP 57 Beard et al. (2007)* ------Paukkaungia parva NMMP 54 Beard et al. (2007)* - - - 3.846 1.664 - - Paukkaungia parva - - - 3.846 1.664 7.788 1.609 Area = MD*BL, shape = MD/BL, *Indicates additional novel measurements were recorded from published photos using ImageJ method described above, **Indicates composite specimens, ***Indicates a species was synonymized with another genus/species. Population averages in bold. All measurements in millimeters.

74 Table 7: Comparative absolute measurements of upper molars for all taxa sampled in analysis.

Taxon Specimen Reference M1 MD M1 BL M1 Ht. M2 MD M2 BL M3 MD M3 BL

Gingerich et al. Altanius orlovi PSS 20-61 0.98 1.69 0.81 0.98 1.8 0.98 1.47 (1991) Gingerich et al. Altanius orlovi PSS 20-168 1.11 1.72 1 1.07 1.92 0.87 1.62 (1991) Altanius orlovi 1.045 1.705 0.905 1.025 1.86 0.925 1.545 Purgatorius titusi UM 10023 Bukley (1997) 1.9 2.6 - - - - - Purgatorius titusi 90166 Bukley (1997) - - - 2 2.75 - - Purgatorius titusi 90191 Bukley (1997) - - - - - 1.5 2.2 Van Valen and Purgatorius unio UM VP 1597 - - - 1.811 2.955 - - Sloan (1965) Purgatorius unio 1.9 2.6 - 1.906 2.853 1.500 2.200 IRSNB 56, Teilhardina belgica 58, 59, 1269, Szalay (1976) 1.635 2.617 - 1.671 2.742 1.222 2.115 4325** CM 67854, CM 77210, Teilhardina magnoliana CM 70431, Beard (2008) 1.768 2.681 - 1.862 3.147 - - and CM 70436** Teilhardina asiatica IVPP V12358 Ni et al. (2004) 1.908 2.527 0.78 1.771 2.982 1.085 2.061 Teilhardina 1.770 2.608 0.780 1.768 2.957 1.154 2.088 Cantius torresi UM 83475 Gingerich (1986) 3.304 4.34 1.917 3.323 5.254 2.413 4.384 Cantius torresi UM 95107 Gingerich (1995) 3.535 4.86 1.815 3.426 5.133 2.348 4.27 Cantius ralstoni AMNH 16089 Rose et al. (1994) 3.474 4.532 - 3.52 5.248 2.63 4.074 Cantius spp. 3.438 4.577 1.866 3.423 5.212 2.464 4.243 Donrussellia provincialis RI 229 Godinot (1981) 2.1 2.7 - - - - - Donrussellia provincialis RI 171 Godinot (1981) - - - 2.2 3.3 - - Donrussellia provincialis RI 173 Godinot (1981) - - - 2.2 3.3 - - Donrussellia provincialis RI 247 Godinot (1981) - - - - - 1.9 2.8 Donrussellia provincialis RI 266 Godinot (1981) - - - - - 1.7 2.6

75 Donrussellia provincialis RI 335 Godinot (1981) - - - - - 1.9 2.7 Donrussellia provincialis 2.100 2.700 - 2.200 3.300 1.833 2.700 PW1995/69- Europolemur klatti Franzen (2004) 4.39 4.52 - - - - - LS PW1995/70- Europolemur klatti Franzen (2004) - - - 4.3 4.33 - - LS Europolemur klatti BUX 80.82 Godinot (1988) 4.07 4.76 - - - - - Europolemur klatti BUX 6.15 Godinot (1988) 4.13 4.51 - - - - - Europolemur klatti BUX 6.16 Godinot (1988) 4.43 5.37 - - - - - Europolemur klatti BUX 80.81 Godinot (1988) - - - 4.25 5.2 - - Europolemur klatti Bchs 305 Godinot (1988) - - - 4.18 5.48 - - Europolemur klatti Bchs 315 Godinot (1988) - - - 4.75 5.52 - - Europolemur klatti BUX 80.80 Godinot (1988) - - - - - 3.57 4.31 Europolemur klatti BUX 6.19 Godinot (1988) - - - - - 3.84 4.91 Europolemur klatti BUX 6.20 Godinot (1988) - - - - - 4.06 4.98 Europolemur klatti BUX 6.21 Godinot (1988) - - - - - 3.97 4.85 Europolemur klatti BUX 80.74 Godinot (1988) - - - - - 3.6 4.41 Europolemur klatti BUX 80.75 Godinot (1988) - - - - - 3.8 4.58 Europolemur klatti 4.255 4.790 - 4.370 5.133 3.807 4.673 Periconodon huerzeleri Bchs 222 Godinot (1988) 2.71 3.35 - 2.86 3.89 2.57 3.66 Periconodon huerzeleri Bchs 640 Godinot (1988) - - - 2.82 4.27 2.62 3.89 Periconodon huerzeleri bchs 450 Godinot (1988) 2.82 4.08 - - - - - Periconodon huerzeleri bchs 312 Godinot (1988) 3.08 4.34 - - - - - Periconodon huerzeleri 2.870 3.923 - 2.840 4.080 2.595 3.775 Marcgodinotius indicus GU 43 Rose et al. (2009) 2.2 2.5 1.043 - - - - Marcgodinotius indicus GU 47 Rose et al. (2009) - - - - - 1.7 2.45 Marcgodinotius indicus GU 48 Rose et al. (2009) - - - - - 1.5 2.45 Marcgodinotius indicus GU 538 Rose et al. (2009) - - - - - 1.8 2.4 Marcgodinotius indicus GU 706 Rose et al. (2009) - - - - - 1.55 2.3 Marcgodinotius indicus 2.2 2.5 1.043 - - 1.638 2.4

76 Protoadapis curvicuspidens Louis-85 Gr Russell et al. (1967) 4.6 5.3 - - - - - Protoadapis curvicuspidens AL-5191 Russell et al. (1967) - - - 4.7 5.9 - - Protoadapis curvicuspidens Louis-89 Gr Russell et al. (1967) - - - 5.1 6.5 - - Protoadapis curvicuspidens Louis-91 Gr Russell et al. (1967) - - - 4 5 - - Protoadapis curvicuspidens Louis-102 Gr Russell et al. (1967) - - - 4.5 5.5 - - Protoadapis curvicuspidens Louis-90 Gr Russell et al. (1967) - - - - - 3.9 5.8 Protoadapis curvicuspidens Louis-100 Gr Russell et al. (1967) - - - - - 4.6 5.5 Protoadapis curvicuspidens Louis-107 Gr Russell et al. (1967) - - - - - 3.3 5.5

Protoadapis curvicuspidens 4.6 5.3 - 4.575 5.725 3.933 5.6

Asiadapis cambayensis GU 609 Rose et al. (2009) - - - 2.8 3.8 - - Asiadapis cambayensis GU 610 Rose et al. (2009) - - - - - 2.2 3.35 Asiadapis cambayensis GU 656 Rose et al. (2009) 2.7 3.55 1.258 2.65 4 2 3.2 Asiadapis cambayensis GU 700 Rose et al. (2009) - - - - - 2.3 3.3 Asiadapis cambayensis GU 701 Rose et al. (2009) - - - 2.9 3.8 - - Asiadapis cambayensis GU 702 Rose et al. (2009) 2.6 3.2 - - - - - Asiadapis cambayensis GU 1550 Rose et al. (2009) 2.55 3.3 - - - - - Asiadapis cambayensis 2.617 3.350 1.258 2.783 3.867 2.167 3.283 GSP-UM Gunnell et al. Panobius russelli - - - - - 1.4 2.5 6787 (2008) GSP-UM Gunnell et al. Panobius russelli - - - - - 1.4 2.2 6788 (2008) GSP-UM Gunnell et al. Panobius russelli - - - - - 1.4 2.4 6790 (2008) GSP-UM Gunnell et al. Panobius russelli - - - 2.3 1.3 - - 6793 (2008) GSP-UM Gunnell et al. Panobius russelli - - - 2 1.5 - - 6798 (2008) GSP-UM Gunnell et al. Panobius russelli - - - 2 1.5 - - 6799 (2008) GSP-UM Gunnell et al. Panobius russelli - - - - - 1.1 2.2 6789 (2008)

77 GSP-UM Gunnell et al. Panobius russelli 2.1 2.9 - - - - - 6784 (2008) GSP-UM Gunnell et al. Panobius russelli 1.9 2.9 - - - - - 6785 (2008) GSP-UM Gunnell et al. Panobius russelli 2 3 - - - - - 6786 (2008) Panobius russelli 2 2.933 - 2.100 1.433 1.325 2.325 Unnumbered, Gingerich et al. Huanghonius stehlini - - - 3.9 5.9 - - Uppsala (1994) Huanghonius stehlini - - - 3.9 5.9 - - Gingerich et al. Rencunius zhoui IVPP 5313 4.2 5.3 - - - - - (1994) Rencunius zhoui 4.2 5.3 - - - - - Lushius quilinensis Type Chow (1961) 4.5 6.9 - 4.6 6 - - Lushius quilinensis 4.5 6.9 - 4.6 6 - - Qi and Beard Guangxilemur tongi IVPP 11652 - - - 6.6 9.1 - - (1998) Guangxilemur tongi - - - 6.6 9.1 - - Marivaux et al. Guangxilemur singsilai DBC 2167 5.173 6.615 - - - - - (2002) Marivaux et al. Guangxilemur singsilai DBC 2168 - - - 5.049 6.885 - - (2002) Guanxilemur singsilai 5.173 6.615 - 5.049 6.885 - - MD = maximum mesiodistal length, BL = maximum buccal-lingual width, Ht. = crown height at paracone, *Indicates additional novel measurements were recorded from published photos using ImageJ method described above, **Indicates composite specimens, ***Indicates a species was synonymized with another genus/species. Population averages in bold. All measurements in millimeter

78 Table 8: Comparative absolute measurements of upper premolars for all taxa sampled in analysis

Taxon Specimen Reference P3 MD P3 BL P3 Ht. P4 MD P4 BL P4 Ht.

Altanius orlovi PSS 20-61 Gingerich et al. (1991) 1 1.26 0.96 0.96 1.37 1.08 Altanius orlovi PSS 20-168 Gingerich et al. (1991) - - - 0.95 1.62 1.07 Altanius orlovi 1 1.26 0.96 0.955 1.495 1.075 IRSNB 56, Teilhardina belgica 58, 59, 1269, Szalay (1976) 1.389 2.027 - 1.536 2.233 - 4325** CM 67854, CM 77210, Teilhardina magnoliana CM 70431, Beard (2008) 1.111 1.57 - 1.535 2.303 - and CM 70436** Teilhardina asiatica IVPP V12358 Ni et al. (2004) 1.615 2.003 1.181 1.638 2.412 1.191 Teilhardina 1.372 1.867 1.181 1.570 2.316 1.191 Cantius torresi UM 95107 Gingerich (1995) - - - 3.198 4.107 - Cantius torresi UM 95796 Gingerich (1995) - - - 2.78 3.47 - AMNH Cantius ralstoni Rose et al. (1994) - - - 2.833 3.634 - 16089 Cantius spp. - - - 2.937 3.737 - Donrussellia provincialis RI 160 Godinot (1981) - - - 2 2.4 - Donrussellia provincialis - - - 2.000 2.400 - Europolemur klatti BUX 80.83 Godinot (1988) - - - 3.27 3.91 - Europolemur klatti BUX 80.79 Godinot (1988) 3.64 3.74 - - - - Europolemur klatti BUX 66.146 Godinot (1988) 3.71 3.85 - - - - Europolemur klatti BUX 80.77 Godinot (1988) - - - 3.48 4.12 - Europolemur klatti 3.675 3.795 - 3.375 4.015 - Asiadapis cambayensis GU 34 Rose et al. (2009) - - - 2.55 2.95 - Asiadapis cambayensis GU 330 Rose et al. (2009) 2.3 2.75 1.421 - - - Asiadapis cambayensis GU 727 Rose et al. (2009) - - - 2.407 3.008 -

79 Asiadapis cambayensis 2.3 2.75 1.421 2.4785 2.979 - Rencunius IVPP 5313 Gingerich et al. (1994) - - - 2.97 4.47 - Rencunius zhoui - - - 2.97 4.47 - Lushius quilinensis Type Chow (1961) - - - 3.7 4.9 - Lushius quilinensis - - - 3.7 4.9 - Guangxilemur singsilai DBC 2166 Marivaux et al. (2002) - - - 4.347 5.907 - Guangxilemur singsilai - - - 4.347 5.907 - MD = maximum mesiodistal length, BL = maximum buccal-lingual width, Ht. = crown height at paracone, *Indicates additional novel measurements were recorded from published photos using ImageJ method described above, **Indicates composite specimens. Population averages in bold. All measurements in millimeters.

80 Table 9: Comparative shape and area values of upper molars derived from absolute lower molar measurements.

Taxon Specimen Reference M1 area M1 shape M2 area M2 shape M3 area M3 shape

Gingerich et al. Altanius orlovi PSS 20-61 1.656 0.580 1.764 0.544 1.441 0.667 (1991) Gingerich et al. Altanius orlovi PSS 20-168 1.909 0.645 2.054 0.557 1.409 0.537 (1991) Altanius orlovi 1.783 0.613 1.909 0.551 1.425 0.602 Purgatorius titusi UM 10023 Bukley (1997) 4.940 0.731 - - - - Purgatorius titusi 90166 Bukley (1997) - - 5.500 0.727 - - Purgatorius titusi 90191 Bukley (1997) - - - - 3.300 0.682 Van Valen and Purgatorius unio UM VP 1597 - - 5.352 0.613 - - Sloan (1965) Purgatorius unio 4.940 0.731 5.426 0.670 3.300 0.682 IRSNB 56, 58, 59, Teilhardina belgica Szalay (1976) 4.279 0.625 4.582 0.609 2.585 0.578 1269, 4325**

CM 67854, CM 77210, CM Teilhardina magnoliana Beard (2008) 4.740 0.659 5.860 0.592 - - 70431, CM 70436**

Teilhardina asiatica IVPP V12358 Ni et al. (2004) 4.822 0.755 5.281 0.594 2.236 0.526 Teilhardina 4.613 0.680 5.241 0.598 2.410 0.552 Cantius torresi UM 83475 Gingerich (1986) 14.339 0.761 17.459 0.632 10.579 0.550 Cantius torresi UM 95107 Gingerich (1995) 17.180 0.727 17.586 0.667 10.026 0.550 Cantius ralstoni AMNH 16089 Rose et al. (1994) 15.744 0.767 18.473 0.671 10.715 0.646 Cantius spp. 15.755 0.752 17.839 0.657 10.440 0.582 Donrussellia provincialis RI 229 Godinot (1981) 5.670 0.778 - - - - Donrussellia provincialis RI 171 Godinot (1981) - - 7.260 0.667 - - Donrussellia provincialis RI 173 Godinot (1981) - - 7.260 0.667 - - Donrussellia provincialis RI 247 Godinot (1981) - - - - 5.320 0.679

81 Donrussellia provincialis RI 266 Godinot (1981) - - - - 4.420 0.654 Donrussellia provincialis RI 335 Godinot (1981) - - - - 5.130 0.704 Donrussellia provincialis 5.670 0.778 7.260 0.667 4.957 0.679 Europolemur klatti PW1995/69-LS Franzen (2004) 19.843 0.971 - - - - Europolemur klatti PW1995/70-LS Franzen (2004) - - 18.619 0.993 - - Europolemur klatti BUX 80.82 Godinot (1988) 19.373 0.855 - - - - Europolemur klatti BUX 6.15 Godinot (1988) 18.626 0.916 - - - - Europolemur klatti BUX 6.16 Godinot (1988) 23.789 0.825 - - - - Europolemur klatti BUX 80.81 Godinot (1988) - - 22.100 0.817 - - Europolemur klatti Bchs 305 Godinot (1988) - - 22.906 0.763 - - Europolemur klatti Bchs 315 Godinot (1988) - - 26.220 0.861 - - Europolemur klatti BUX 80.80 Godinot (1988) - - - - 15.387 0.828 Europolemur klatti BUX 6.19 Godinot (1988) - - - - 18.854 0.782 Europolemur klatti BUX 6.20 Godinot (1988) - - - - 20.219 0.815 Europolemur klatti BUX 6.21 Godinot (1988) - - - - 19.255 0.819 Europolemur klatti BUX 80.74 Godinot (1988) - - - - 15.876 0.816 Europolemur klatti BUX 80.75 Godinot (1988) - - - - 17.404 0.830 Europolemur klatti 20.408 0.892 22.461 0.858 17.832 0.815 Periconodon huerzeleri Bchs 222 Godinot (1988) 9.079 0.809 11.125 0.735 9.406 0.702 Periconodon huerzeleri Bchs 640 Godinot (1988) - - 12.041 0.660 10.192 0.674 Periconodon huerzeleri bchs 450 Godinot (1988) 11.506 0.691 - - - - Periconodon huerzeleri bchs 312 Godinot (1988) 13.367 0.710 - - - - Periconodon huerzeleri 11.317 0.737 11.583 0.698 9.799 0.688 Marcgodinotius indicus GU 47 Rose et al. (2009) - - - - 4.165 0.694 Marcgodinotius indicus GU 48 Rose et al. (2009) - - - - 3.675 0.612 Marcgodinotius indicus GU 538 Rose et al. (2009) - - - - 4.32 0.750 Marcgodinotius indicus GU 706 Rose et al. (2009) - - - - 3.565 0.674 Marcgodinotius indicus 5.5 0.88 - - 3.931 0.683 Russell et al. Protoadapis curvicuspidens Louis-85 Gr 24.38 0.868 - - - - (1967)

82 Russell et al. Protoadapis curvicuspidens AL-5191 - - 27.73 0.797 - - (1967) Russell et al. Protoadapis curvicuspidens Louis-89 Gr - - 33.15 0.785 - - (1967) Russell et al. Protoadapis curvicuspidens Louis-91 Gr - - 20 0.800 - - (1967) Russell et al. Protoadapis curvicuspidens Louis-102 Gr - - 24.75 0.818 - - (1967) Russell et al. Protoadapis curvicuspidens Louis-90 Gr - - - - 22.62 0.672 (1967) Russell et al. Protoadapis curvicuspidens Louis-100 Gr - - - - 25.3 0.836 (1967) Russell et al. Protoadapis curvicuspidens Louis-107 Gr - - - - 18.15 0.600 (1967)

Protoadapis curvicuspidens 24.38 0.868 26.408 0.800 22.023 0.703

Asiadapis cambayensis GU 609 Rose et al. (2009) - - 10.64 0.737 - - Asiadapis cambayensis GU 610 Rose et al. (2009) - - - - 7.37 0.657 Asiadapis cambayensis GU 656 Rose et al. (2009) 9.585 0.761 10.6 0.663 6.4 0.625 Asiadapis cambayensis GU 700 Rose et al. (2009) - - - - 7.59 0.697 Asiadapis cambayensis GU 701 Rose et al. (2009) - - 11.02 0.763 - - Asiadapis cambayensis GU 702 Rose et al. (2009) 8.32 0.813 - - - - Asiadapis cambayensis GU 1550 Rose et al. (2009) 8.415 0.773 - - - - Asiadapis cambayensis 8.773 0.782 10.753 0.721 7.120 0.660 Gunnell et al. Panobius russelli GSP-UM 6787 - - - - 3.5 0.560 (2008) Gunnell et al. Panobius russelli GSP-UM 6788 - - - - 3.08 0.636 (2008) Gunnell et al. Panobius russelli GSP-UM 6790 - - - - 3.36 0.583 (2008) Gunnell et al. Panobius russelli GSP-UM 6793 - - 2.99 1.769 - - (2008) Gunnell et al. Panobius russelli GSP-UM 6798 - - 3 1.333 - - (2008) Gunnell et al. Panobius russelli GSP-UM 6799 - - 3 1.333 - - (2008)

83 Gunnell et al. Panobius russelli GSP-UM 6789 - - - - 2.42 0.5 (2008) Gunnell et al. Panobius russelli GSP-UM 6784 6.09 0.724 - - - - (2008) Gunnell et al. Panobius russelli GSP-UM 6785 5.51 0.655 - - - - (2008) Gunnell et al. Panobius russelli GSP-UM 6786 6 0.667 - - - - (2008) Panobius russelli 5.867 0.682 2.997 1.479 3.090 0.570 Unnumbered, Gingerich et al. Huanghonius stehlini - - 23.01 0.661 - - Uppsala (1994) Huanghonius stehlini - - 23.01 0.661 - - Gingerich et al. Rencunius zhoui IVPP 5313 22.26 0.792 - - - - (1994) Rencunius zhoui 22.26 0.792 - - - - Lushius quilinensis Type Chow (1961) 31.05 0.652 27.6 0.767 - 0.75 Lushius quilinensis 31.05 0.652 27.6 0.767 - 0.75 Qi and Beard Guangxilemur tongi IVPP 11652 - - 60.06 0.725 - - (1998) Guangxilemur tongi - - 60.06 0.725 - - Marivaux et al. Guangxilemur singsilai DBC 2167 34.219 0.782 - - - - (2002) Marivaux et al. Guangxilemur singsilai DBC 2168 - - 34.762 0.733 - - (2002) Guangxilemur singsilai 34.219 0.782 34.762 0.733 - - Area = MD*BL, shape = MD/BL, *Indicates additional novel measurements were recorded from published photos using ImageJ method described above, **Indicates composite specimens. Population averages in bold. All measurements in millimeters.

84 Table 10: Comparative shape and area values of upper premolars derived from absolute lower molar measurements

P1 P1 P2 P2 P3 P3 P4 P4 Taxon Specimen Reference area shape area shape area shape area shape

Gingerich et al. Altanius orlovi PSS 20-61 - - - - 1.26 0.794 1.315 0.701 (1991) Gingerich et al. Altanius orlovi PSS 20-168 ------1.539 0.586 (1991) Altanius orlovi - - - - 1.26 0.794 1.427 0.644 IRSNB 56, 58, 59, Teilhardina belgica Szalay (1976) - - - - 2.816 0.685 3.430 0.688 1269, 4325**

CM 67854, CM 77210, CM Teilhardina magnoliana Beard (2008) - - - - 1.744 0.708 3.535 0.667 70431, CM 70436**

Teilhardina asiatica IVPP V12358 Ni et al. (2004) 0.267 0.821 0.358 1.632 3.235 0.806 3.951 0.679 Teilhardina 0.267 0.821 0.358 1.632 2.598 0.733 3.639 0.678 Cantius torresi UM 95107 Gingerich (1995) ------13.134 0.779 Cantius torresi UM 95796 Gingerich (1995) ------9.647 0.801 Cantius ralstoni AMNH 16089 Rose et al. (1994) ------10.295 0.780 Cantius spp. ------11.025 0.786 Donrussellia provincialis RI 160 Godinot (1981) ------4.800 0.833 Donrussellia provincialis ------4.800 0.833 Europolemur klatti BUX 80.85 Godinot (1988) - - 2.247 2.000 - - - - Europolemur klatti BUX 80.83 Godinot (1988) ------12.786 0.836 Europolemur klatti BUX 80.79 Godinot (1988) - - - - 13.614 0.973 - - Europolemur klatti BUX 66.146 Godinot (1988) - - - - 14.284 0.964 - - Europolemur klatti BUX 80.77 Godinot (1988) ------14.338 0.845 Europolemur klatti - - 2.247 2.000 13.949 0.968 13.562 0.840 Asiadapis cambayensis GU 34 Rose et al. (2009) ------7.523 0.864 Asiadapis cambayensis GU 330 Rose et al. (2009) - - - - 6.325 0.836 - -

85 Asiadapis cambayensis GU 727 Rose et al. (2009) ------0.800 Asiadapis cambayensis - - - - 6.325 0.836 7.523 0.832 Gingerich et al. Rencunius zhoui IVPP 5313 ------13.276 0.664 (1994) Rencunius zhoui ------13.276 0.664 Lushiu quilinensis Type Chow (1961) ------18.13 0.755 Lushius quilinensis ------18.13 0.755 Marivaux et al. Guangxilemur singsilai DBC 2166 ------25.678 0.736 (2002) Guangxilemur singsilai ------25.678 0.736 Area = MD*BL, shape = MD/BL, *Indicates additional novel measurements were recorded from published photos using ImageJ method described above, **Indicates composite specimens. Population averages in bold. All measurements in millimeters.

86 Table 11: “Conservative” comparative relative measurements of the lower dentition of all taxa sampled P1 P2 P3 P3 P3 P4 P4 M1 M1 M2 P2 P3 Ht. area / area / MD / MD / Ht. / MD / Ht. / area area / area / Taxon Specimen Reference Ht. / / M1 P2 P3 P4 M1 P4 M1 M1 / M2 M3 M3 P3 Ht. Ht. area area MD MD Ht. MD Ht. area area area Dashzeveg and GISPS McKenna Altanius orlovi coll.7, No. ------0.833 1.069 0.927 0.848 0.915 (1977); Rose and 20-8 Krause (1984)* Gingerich et al. Altanius orlovi PSS 20-58 - - - 1.043 0.850 0.979 1.051 0.814 1.073 0.888 0.757 0.852 (1991)* Gingerich et al. Altanius orlovi PSS 20-85 ------0.823 (1991) PSS 20- Gingerich et al. Altanius orlovi - - - 0.830 0.690 1.024 1.132 0.832 1.105 0.838 - - 136 (1991)* Altanius - - - 0.937 0.770 1.002 1.091 0.826 1.082 0.884 0.802 0.864 Purgatorius UCMP Clemens (1974)* 0.705 0.909 0.757 0.630 0.693 0.836 0.833 1.207 1.976 1.172 0.593 unio 107406 Purgatorius - 0.705 0.909 0.757 0.630 0.693 0.836 0.833 1.207 1.976 1.172 0.593 Szalay (1976); Teilhardina IRSNB Smith et al. - - - 0.980 0.739 0.845 1.070 0.754 1.267 0.972 1.243 1.279 belgica M64 (2006)* CM 73229, CM Teilhardina 67856, Beard (2008)* ------0.645 - 1.305 1.160 0.889 magnoliana CM 70435, and CM 70427** Teilhardina IVPP Ni et al. (2004)* 0.556 0.321 0.530 0.973 0.738 0.820 1.360 0.758 1.659 1.026 1.536 1.497 asiatica V12358 Teilhardina 0.556 0.321 0.530 0.977 0.738 0.832 1.215 0.719 1.463 1.101 1.313 1.221 UM Gingerich Cantius torresi - - - 0.840 0.708 - 1.134 0.843 - - - - 83470 (1986)* UM Gingerich Cantius torresi 0.610 0.445 0.809 0.766 0.671 0.856 0.947 0.876 1.106 0.915 0.940 1.028 101958 (1995)*

87 UM Gingerich Cantius torresi - 0.508 - 0.860 0.723 0.905 - 0.841 - 0.870 - 87341 (1995)* UM Cantius torresi Gingerich (1995) ------0.935 87341 Cantius torresi 0.610 0.477 0.809 0.822 0.701 0.881 1.041 0.853 1.106 0.893 0.940 0.982 YPM Rose et al. Cantius ralstoni - - - 0.852 0.784 1.028 1.248 0.921 1.214 0.846 0.874 1.034 23317 (1994)* Rose et al. Cantius ralstoni UW 8842 - - - 0.891 0.829 0.845 1.059 0.930 1.254 0.867 - - (2011)* Cantius ralstoni - - - 0.871 0.806 0.936 1.153 0.925 1.233 0.856 0.874 1.033 Donrussellia RI 170 Godinot (1981)* ------0.815 0.734 0.901 provincialis Donrussellia RI 430 Godinot (2015)* - - - 0.866 0.729 0.986 1.257 0.842 1.274 - - - provincialis Donrussellia - - - 0.866 0.729 0.986 1.256 0.841 1.274 0.815 0.734 0.900 provincialis Europolemur Bchs 247 Godinot (1988)* - 0.412 - - 0.835 ------klatti Europolemur BUX Godinot (1988) ------0.920 0.922 1.002 klatti 80.107 Europolemur - 0.412 - - 0.835 - - - - 0.920 0.922 1.002 klatti Periconodon Bchs 494 Godinot (1988) ------0.873 0.720 0.825 huerzeleri Periconodon Bchs 313 Godinot (1988) ------0.854 - - huerzeleri Periconodon Bchs 270 Godinot (1988) ------huerzeleri Periconodon Bchs 495 Godinot (1988)* ------0.899 - 0.989 1.240 1.254 huerzeleri Periconodon ------0.899 - 0.905 0.980 1.040 huerzeleri Femenias-Gaul Agerinia roselli IPS-2543 - - - 0.819 - 0.996 ------et al. (2016)* Femenias-Gaul Agerinia roselli IPS-1981 ------1.004 et al. (2016)* Femenias-Gaul Agerinia roselli IPS-2541 ------1.184 et al. (2016)*

88 Agerinia roselli - - - 0.819 - 0.996 - - - - - 1.094 Protoadapis MNHN Russell et al. ------0.955 curvicuspidens AL-5179 (1967) Protoadapis MNHN Russell et al. - 0.361 - - - - - 1.146 - 0.899 - - curvicuspidens AL-5180 (1967) MNHN- Protoadapis Russell et al. Louis-15 - - - 0.878 0.956 1.050 1.609 1.089 1.533 0.904 0.791 0.875 curvicuspidens (1967)* Ma Protoadapis Lyon Russell et al. ------0.964 - - - - curvicuspidens 1991 (1967) Protoadapis - 0.361 - 0.878 0.956 1.050 1.609 1.066 1.533 0.902 0.791 0.915 curvicuspidens Marcgodinotius Rose et al. GU 227 - - - 0.826 ------1.198 indicus (2009)* Marcgodinotius - - - 0.826 - - - - - 1.198 indicus Asiadapis GU/RSR/ Rose et al. - - - 1.000 0.912 1.098 1.538 0.912 1.401 0.914 - - cambayensis VAS-6 (2007)* Asiadapis Rose et al. GU 745 - - - 0.983 0.952 - - 0.968 1.535 0.917 - - cambayensis (2009)* Suratius Rose et al. IITR 928 - - - 1.055 ------robustus*** (2009) Asiadapis - - - 1.013 0.932 1.098 1.538 0.940 1.468 0.915 - - cambayensis Adapoides IVPP Beard et al. ------0.819 troglodytes V11023 (1994)* Adapoides ------0.819 troglodytes Gilbert et al. Unnumber Hoanghonius (2017); ed, ------1.063 stehlini Gingerich et al Uppsala (1994) Hoanghonius IVPP Tong et al. - 0.516 0.787 0.882 0.769 1.102 1.598 0.872 1.450 0.910 0.848 0.932 stehlini 10220 (1999)* Hoanghonius - 0.516 0.787 0.882 0.769 1.102 1.598 0.872 1.450 0.910 0.848 0.997 stehlini IVPP Gilbert et al. Rencunius zhoui ------0.947 1.125 0.776 - - 5312 (2017);

89 Gingerich et al (1994)

Rencunius ------0.947 1.125 0.776 - - zhoui Gilbert et al. Wailekia TF 2632 (2017); Ducroq ------1.101 orientale et al. (1995)* Wailekia ------1.101 orientale MD = maximum mesiodistal length, Area = MD*BLTal, Ht. = crown height at protoconid, *Indicates additional novel measurements were recorded from published photos using ImageJ method described above, **Indicates composite specimens, ***Indicates a species was synonymized with another genus/species. Population averages in bold. All measurements in millimeters.

90

Table 12: “Liberal” comparative relative measurements of lower dentition using population average dental measurements for all taxa Avg. Avg. P1 P2 Avg. P2 Avg. P3 Avg. P3 Avg. P3 Avg. P4 Avg. M1 Avg. M1 Avg. M2 Avg. P3 Avg. P4 area / area / Ht / MD / MD / Ht / MD / area / area / area / Taxon Ht / Avg. Ht / Avg. Avg. Avg. Avg. P3 Avg. P4 Avg. Mean M1 Avg. M1 Avg. M2 Avg. M3 Avg. M3 P4 Ht M1 Ht P2 P3 Ht MD M1 MD Ht MD area area area area area Altanius - - - 0.913 0.754 0.963 1.040 0.827 1.080 0.923 0.780 0.845 Purgatorius - 0.705 0.909 0.821 0.677 0.881 0.966 0.824 1.095 1.289 1.174 0.911 Teilhardina 0.556 0.360 0.527 1.033 0.743 0.832 1.293 0.720 1.554 1.079 1.308 1.212 Cantius spp. 0.509 0.400 0.709 0.844 0.746 0.906 1.085 0.884 1.198 0.891 0.934 1.048 Donrussellia - - - 0.865 0.765 0.986 1.223 0.884 1.240 0.927 0.962 1.038 provincialis Europolemur - 0.349 - 0.888 0.850 - - 0.957 - 0.831 0.841 1.013 klatti Periconodon ------0.879 n/a 0.965 0.991 1.027 huerzeleri Agerinia roselli - - - 0.819 0.701 0.996 1.262 0.855 1.267 0.949 1.014 1.068 Protoadapis - 0.430 - 0.954 1.029 1.050 1.609 1.078 1.533 0.929 0.877 0.944 curvicuspidens Marcgodinotius - - - 0.882 0.848 1.113 1.496 0.961 1.344 0.948 1.144 1.207 indicus Asiadapis - - - 0.994 0.916 1.161 1.705 0.921 1.469 0.945 1.036 1.096 cambayensis Adapoides ------0.819 troglodytes Panobius ------0.953 - 1.029 1.040 1.011 russelli Hoanghonius - 0.516 0.787 0.882 0.769 1.102 1.598 0.872 1.450 0.867 0.863 0.996 stehlini Rencunius zhoui ------0.947 1.125 0.752 0.874 1.161 Wailekia ------1.101 orientale Paukkaungia - - - 0.715 0.705 0.757 0.679 0.986 0.898 1.103 - - parva

91 Population averages calculated from absolute dental measurements for all specimens sampled (see Tables 3 & 5). MD = maximum mesiodistal length, Area = MD*BLTal, ShapeTri, Ht. = crown height at protoconid. All measurements in millimeters Table 13: “Conservative” comparative relative measurements of upper dentition of all taxa sampled P3 area P3 area P4 area M1 area M1 area M2 area P3 Ht / P4 Ht / Taxon Specimen Reference / P4 / M1 / M1 / M2 / M3 / M3 P4 Ht M1 Ht area area area area area area Gingerich et al. Altanius orlovi PSS 20-61 0.958 0.761 0.889 0.794 1.333 0.939 1.150 1.224 (1991)* Gingerich et al. Altanius orlovi PSS 20-168 - - - 0.806 1.070 0.929 1.355 1.458 (1991)* Altanius 0.958 0.761 0.889 0.800 1.202 0.934 1.252 1.341 IRSNB 56, 58, Teilhardina belgica 59, 1269, Szalay (1976)* 0.821 0.658 - 0.802 - 0.934 1.656 1.773 4325** CM 67854, CM 77210, Teilhardina magnoliana Beard (2008) 0.493 0.368 - 0.746 - 0.809 - - CM 70431, CM 70436** Teilhardina asiatica IVPP V12358 Ni et al. (2004)* 0.819 0.671 0.992 0.819 1.527 0.913 2.156 2.362 Teilhardina 0.711 0.566 0.992 0.789 1.527 0.913 2.156 2.362 Cantius torresi UM 83475 Gingerich (1986) - - - - - 0.821 1.356 1.650 Cantius torresi UM 95107 Gingerich (1995) - - - 0.764 - 0.977 1.714 1.754 Cantius torresi UM 95796 Gingerich (1995) ------Cantius torresi - - - 0.764 - 0.899 1.535 1.702 Cantius ralstoni AMNH 16089 Rose et al. (1994) - - - 0.654 - 0.852 1.469 1.724 Cantius ralstoni - - - 0.654 - 0.852 1.469 1.724 Periconodon huerzeleri Bchs 222 Godinot (1988) - - - - - 0.816 0.965 1.183 Periconodon huerzeleri Bchs 640 Godinot (1988) ------1.181 Periconodon huerzeleri - - - - - 0.816 0.965 1.182 Asiadapis cambayensis GU 656 Rose et al. (2009) - - - - - 0.904 1.498 1.656 Asiadapis cambayensis - - - - - 0.904 1.498 1.656 Gingerich et al. Rencunius zhoui IVPP 5313 - - - 0.596 - - - - (1994) Rencunius zhoui - - - 0.596 - Lushius quilinensis Type Chow (1961) - - - 0.584 - 1.125 2.682 2.331 Lushius quilinensis - - - 0.584 - 1.125 2.682 2.331

92 MD = maximum mesiodistal length, Area = MD*BL, Ht. = crown height at paracone, *Indicates additional novel measurements were recorded from published photos using ImageJ method described above, **Indicates composite specimens. Population averages in bold. All measurements in millimeters. Table 14: “Liberal” comparative relative measurements using population average dental measurements for all taxa sampled Avg. P3 area Avg. P3 area Avg. M2 area Avg. P3 Ht / Avg. P4 area / Avg. P4 Ht / Avg. M1 area / Avg. M1 area Taxon / Avg. P4 / Avg. M1 / Avg. M3 Avg. P4 Ht Avg. M1 area Avg. M1 Ht Avg. M2 area / Avg. M3 area area area area Altanius orlovi 0.883 0.707 0.893 0.801 1.188 0.934 1.251 1.340 Purgatorius unio - - - - - 0.910 1.497 1.644 Teilhardina 0.714 0.563 0.992 0.789 1.527 0.880 1.914 2.174 Cantius spp. - - - 0.700 - 0.883 1.509 1.709 Donrussellia - - - 0.847 - 0.781 1.144 1.465 provincialis Europolemur 1.029 0.683 - 0.665 - 0.909 1.144 1.260 klatti Periconodon - - - - - 0.977 1.155 1.182 huerzeleri Marcgodinotius ------1.399 - indicus Protoadapis - - - - - 0.923 1.107 1.199 curvicuspidens Asiadapis 0.841 0.721 - 0.857 - 0.816 1.232 1.510 cambayensis Panobius russelli - - - - - 1.958 1.899 0.970 Huanghonius ------stehlini Rencunius zhoui - - - 0.596 - Lushius - - - 0.584 - 1.125 2.682 2.331 quilinensis Guangxilemur ------tongi Guanxilemur - - - 0.750 - 0.984 - - singsilai Population averages calculated from absolute dental measurements for all specimens sampled (see Tables 7 & 8) Area = MD*BL, Ht. = crown height at paracone. All measurements in millimeters.

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