# Maxilla of Siamopithecus eocaenus (Anthropoidea, Primates) from the Paleogene of Krabi, Thailand, and its taxonomic status

Yaowalak Chaimanee, Sasa-On Khansubha, Olivier Chavasseau, Arnaud Mazurier, Jean-Jacques Jaeger

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Yaowalak Chaimanee, Sasa-On Khansubha, Olivier Chavasseau, Arnaud Mazurier, Jean-Jacques Jaeger. Maxilla of Siamopithecus eocaenus (Anthropoidea, Primates) from the Paleogene of Krabi, Thailand, and its taxonomic status. Journal of Human Evolution, 2025, 198, pp.103614. 10.1016/j.jhevol.2024.103614. hal-04911167

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Maxilla of *Siamopithecus eocaenus* (Anthropoidea, Primates) from the Paleogene of
Krabi, Thailand, and its taxonomic status

Yaowalak CHAIMANEEa,*, Sasa-On KHANSUBHAb, Olivier CHAVASSEAUa, Arnaud MAZURIERc, Jean-Jacques JAEGERa

<sup>a</sup>Laboratory Paleontology, Evolution, Paleoecosystems and Paleoprimatology
)PALEVOPRIM), UMR 7262 CNRS, University of Poitiers, France.

<sup>b</sup>Department of Mineral Resources, Rama VI Road, Bangkok 10400, Thailand.

C Institute of Chemistry of Environments and Materials of Poitiers (IC2MP, UMR 7285 CNRS, University of Poitiers, France.

*Corresponding author*: E-mail address: Y. Chaimanee [yao.chaimanee@univ-poitiers.fr](mailto:yao.chaimanee@univ-poitiers.fr)

**Keywords:** Amphipithecidae, Anthropoid, Catarrhine, Primates, Phylogenetic, Southeast Asia

Chaimanee Y, Khansubha S, Chavasseau O, Mazurier A, Jaeger J-J. 2025. Maxilla of
*Siamopithecus eocaenus* (Anthropoid, Primates) from the Paleogene of Krabi, Thailand, and
its taxonomic status. Journal of Human Evolution 198 (103614):

DOI:10.1016/j.jhevol.2024.103614
Abstract

Detailed descriptions of the maxillae of *Siamopithecus eocaenus*, discovered from the latest Eocene/earliest Oligocene lignite mine in the Krabi basin of Peninsular Thailand are presented. They include the morphology of P³–M³, the palate, a partial orbital region, and the zygomatic root. The specimen exhibits distinctive dental features including a single-rooted P² alveolus, a protocone on P³ and P⁴, and a true hypocone on upper molars, indicating its derived anthropoid dentition. Comparative studies and virtual reconstructions of the facial anatomy reveal close affinities with diurnal anthropoids rather than the notharctid strepsirrhines. The reconstructed facial morphology of *Siamopithecus* displays a short and sub-vertically oriented face, significant orbital convergence (72.1°), and frontation (81.6°), distinguishing it from both fossil and extant strepsirrhines. Moreover, the presence of a thin bony lamina extending from the distal part of the upper preserved area of the zygomatic suggests partial or complete postorbital closure. Phylogenetic analyses suggest an affiliation with amphipithecids, but recent morphological observations challenge this, leading to the proposal of an elevated family-group ranking, Siamopithecidae. Comparison with the most primitive known Afro-Arabian propliopithecid, the Taqah propliopithecid from Oman, reveals similarities in their molar structure but differences in dental formula (retention of P2) and premolar structure. The abrupt appearance of propliopithecids in the early Oligocene of Afro-Arabia, without a local ancestor, contributes to the debate on catarrhine origins, whether from Asia or Africa. However alternative views, based on sister-group relationships with oligopithecids, support an African origin of propliopithecids from an undocumented Afro-Arabian region. This research provides new insights into the evolutionary history of early anthropoids, suggesting a complex biogeographical scenario involving both Asian and African lineages.
# 1. Introduction

*Siamopithecus eocaenus*, described by Chaimanee et al. (1997), is a large-bodied primate known exclusively from the Krabi coal mine of Peninsular Thailand, dating from the latest Eocene to the earliest Oligocene (Benammi et al., 2001). Initially, it was considered closely related to late Eocene anthropoids found in the Pondaung Formation of Myanmar, such as *Pondaungia*, *Ganlea*, and *Myanmarmpithecus*, as part of the Southeast Asian Amphipithecidae (Jaeger et al., 1998, 2020; Ducrocq, 1999; Chaimanee, 2004; Marivaux et al., 2005; Beard et al., 2009; Seiffert, 2012; Chaimanee et al., 2013). However, subsequent discoveries and detailed studies have led to a reclassification of this family together with *Aseanpithecus* (Jaeger et al., 2019) into three subfamilies: Amphipithecinae (including *Pondaungia*, *Ganlea*, and *Myanmarmpithecus*), Aseanpithecinae (*Aseanpithecus*), and *Siamopithecinae* (*Siamopithecus*) (Chaimanee et al., 2024).

The initial remains of *Siamopithecus* including a maxillary fragment bearing P³–M³ (TF 3635) and a mandible fragment preserving the distal part of M₁, and M₂–M₃ (TF 3634), were described by Chaimanee et al. (1997), with subsequent studies providing further details on these specimens by Ducrocq (1999) and Chaimanee (2004). Additional discoveries include a nearly complete right mandible (TF 7624; Chaimanee et al., 2000) found in occlusion with the maxillofacial components (TF 7625). The mandible exhibits several anthropoid characters, such as a vertical symphysis and an oblique alignment of P₃ and P₄ roots. Distinctive characters also include a significant reduction of the single-rooted P₂, an elevated corpus, and the presence of a metaconid on the lingual side of P₄ (Chaimanee et al. 2000; Chaimanee, 2004).

Virtual reconstruction analysis of the maxillofacial remains (TF 7625) conducted by Tafforeau (2004), followed by Zollikofer et al. (2009), revealed a short face and highly
convergent and frontated orbits, suggesting greater morphological resemblance to anthropoids
than to any known strepsirrhines. Other cranial characters, such as the participation of the
maxilla in the lower orbit margin and the absence of an external lacrimal bone, also confirmed
its anthropoid affinities.

Nevertheless, several researchers have raised doubts regarding the anthropoid status of
*Siamopithecus* (Godinot, 1998, 2015; Gunnell and Miller, 2001; Kirk and Simons, 2001;
Gunnell et al., 2002; Kay et al., 2004). Rosenberger et al. (2011) contested the robustness of
the dendrograms presented by Zollikofer et al. (2009), arguing that their broader topology
contained errors and was inconsistent with well-established phylogenies. However, the
phylogenetic status of amphipithecids remained still controversial due to the incompleteness
and non-associated dental and postcranial remains (Ciochon et al., 2001). Some authors
attributed them to adapiforms and interpreted their shared dental characters as resembling
those of anthropoids through homoplasy (Ciochon and Holoyrd, 1994; Ciochon et al. 2001;
Gunnell et al., 2002). Nevertheless, the subsequent discovery of cranial, dental, and
postcranial remains of *Ganlea megacanina* (Dagosto et al., 2010; Marivaux et al., 2010;
Jaeger et al., 2020), demonstrated that *Ganlea*, along with closely related amphipithecine
genera, *Pondaungia* and *Myanmarpithecus*, represent stem anthropoids. The cranial anatomy
of *Ganlea* displays several anthropoid characters, including fused frontal bones, participation
of the maxilla in the lower orbit rim, and an internal lacrimal bone with a sub-vertical lacrimal
canal, although *Ganlea* apparently lacks postorbital closure. This phylogenetic interpretation
was extended to all Myanmar amphipithecines due to their close phylogenetic relationships.
The discovery of the *Ganlea* cranial remain allowed for a precise comparison with
*Siamopithecus*, suggesting that in addition to numerous shared dental and cranial anthropoid
characters (as presented in Table 3 of Zollikofer et al., 2009), *Siamopithecus* differs from
*Ganlea* in most likely possessing partial or complete postorbital closure (Chaimanee et al.,
2024). This, along with differences in dental characters from the Myanmar amphipithecines, raises new questions regarding its phylogenetic position within anthropoid primates and its precise affinities with the Myanmar amphipithecines.

This study aims to provide a comprehensive description and comparative analysis of *Siamopithecus maxillae*, shedding light on its phylogenetic relationships within Asian anthropoid primates and contributing to our understanding of primate diversity during the Paleogene.

## 2. Materials and methods

### 2.1. Locality and age

The fossil described here (TF 7625) was recovered from the main lignite bed of the Bang Mark pit, the same layer as the holotype (TF 3635; Chaimanee et al., 1997). These coal pits were developed by the Electricity Generating Authority of Thailand (EGAT), and extend along a 5 km long NNW-SSE axis (Udomkan et al., 2003). Numerous fossils, including large mammals, turtles, and crocodile remains have been discovered from the main lignite bed of the Bang Mark, Wai Lek, and Bang Pu Dam pits (Chaimanee, 2004; Ducrocq et al., 2021) (Fig. 1). The fossil assemblages found in these three coal pits are considered contemporaneous (Ducrocq et al., 1995, 2021). The composition of this fauna and the reverse magnetic polarity (Chron 13r) observed within the fossiliferous sequence indicate an age range spanning from the latest Eocene to the earliest Oligocene, as Chron C13r corresponds to the Eocene/Oligocene transition (EOT) occurring between 35-33 Ma (Benammi et al., 2001). These diverse faunas inhabited a forested area under a tropical climate (Ducrocq et al. 1995).

### 2.2. Institutional abbreviations

DPC = Duke Primate Center; NMMP = National Museum of Myanmar-Primates; TF = Thai Fossil, the Department of Mineral Resources, Thailand; TQ = Taqah, Oman.
## 2.3. *Measurements*

Dental measurements were taken on the 3D digital models, using Avizo 8. Measurements of mesiodistal length (MD) correspond to the maximum values, and buccolingual width (BL) was measured perpendicular to mesiodistal length taken on upper premolars and molars (Table 1).

## 2.4. *Micro-computed tomography scan imaging*

TF 7625 was scanned using an RX-solutions EasyTom XL Duo microtomograph (platform PLATINA of IC2MP, Université de Poitiers, France), which combines a sealed microfocus x-ray source with a flat panel detector. The scanning was conducted according to the following parameters: 4320 projections over 3 turns, a frame rate of 11.5 fps, a frame averaging of 20, a voltage of 90kV, and a current of 240 µA. Reconstruction of the 16-bit TIFF slices stack was performed with the XAct software package (RX-solutions) at a voxel size of 25.25µm. Surface visualization and manipulation were performed with Avizo 3D 2023.2 (Thermo Fisher Scientific, Waltham, Massachusetts, USA) (Figs. 2, 3).

2.5. *Estimation of orbit size*

The orbit diameter of TF 7625 was estimated using the methodology of Heesy and Ross (2001). This methodology involves using three points positioned along the orbit plane (X, Y, Z) to calculate the orbit radius. The original methodology suggests selecting points along the orbit by using the inferior-most point of the orbit (Y) and two other points (X and Z) equally distant from Y. However, due to limitations in accessing the preserved portion of the orbit in TF 7625, this positioning of points was not feasible. Instead, X and Z were placed at the two extremities of the preserved rim to maximize the length of the orbit rim utilized. The selection of points and the extraction of their coordinates were performed using Avizo 8 software
(ThermoFisher Scientific), which rendered a surface of the maxilla generated from CT-scan
data. The formula used to calculate the orbit diameter (D) was D = 2OY = (WY²+WZ²)/WY,
where O represents the center of the orbit, and W is the midpoint of the XZ segment.

2.6. *Body weight estimation*

We have estimated the body weight of *Siamopithecus* based on the maxilla TF 7625.
We used M², due to the variability observed in M¹. We applied the regression equations
provided by Egi et al. (2004), which were based on the tooth crown area (mesiodistal length,
MD, multiplied by buccolingual width, BL). The calculation for body mass estimated was
derived from anthropoid models.

2.7. *Phylogenetic analysis*

The phylogenetic position of *Siamopithecus* was analyzed using PAUP 4.3.99.169.0
(Swofford, 2003). Three maximum parsimony analyses with heuristic search were performed
using three different taxon-character matrices based on those of Seiffert et al. (2018), Jaeger
et al. (2019), and Jaeger et al. (2020). For full methodological details, see the text and data
matrices in the Supplemental Online Material.

**3. Systematic Paleontology**

**Class Mammalia Linnaeus, 1758**

Order Primates Linnaeus, 1758

Suborder Haplorhini Pocock, 1918

Infraorder Anthropoidea Mivart, 1864

Family Siamopithecidae Beard et al., 2009 (new rank)

Genus *Siamopithecus* Chaimanee et al., 1997
*Type species* *Siamopithecus eocaenus* Chaimanee et al., 1997

*Siamopithecus eocaenus*

**3.1. Description**

TF 7625 consists of both the left and right maxillae, bearing teeth from P³–M³ (Figs. 2, 3), found associated with the right lower jaw (TF 7624) in occlusion. The left maxilla fragment is more complete and includes a part of the zygomatic bone. It bears the left partial orbital region, the root of the zygomatic arch, a segment of the palate, and the canine and P² alveoli (Fig. 2). The premolars and molars on the left maxilla exhibit heavy wear (Fig. 2D), more pronounced than those observed on the right maxilla, indicating potential asymmetrical chewing patterns (Tafforeau, 2004). The orbital floor, formed by the zygomatic and frontal processes of the maxilla, is also discernible. Post-fossilization damage is evident along the zygomaxillary suture. The palatine is incomplete, missing both proximal and distal parts (Fig. 2D). The right maxilla is less complete and slightly deformed (Fig. 3). The zygomatic bone is broken at its base. Its canine and P² alveoli are also present, and its premolars and molars exhibit less wear than those on the left. In lateral view, the lower part of the premaxilla/maxilla suture is visible and vertically oriented. Although it could be interpreted as a crack, its position and orientation in front of the canine strongly suggest that it corresponds to the premaxilla/maxilla suture. Similar to the left maxilla, the right maxilla also displays the orbital floor. Both fragments display a thick maxillary bone in the vertical section (~14.5 mm), consisting of alveolar bone containing long premolars and molar roots. The dental arcades are curved and parabolic in shape (Zollikofer et al., 2009). The enamel of the teeth
appears relatively thin and prismatic, lacking Hunter-Schreger bands (Tafforeau, 2004).

Detailed measurements are provided in Tables 1 and 2.

**Dentition** TF 7625 preserves P³–M³ on both the left and right sides (Figs. 2D, 3D). The occlusal surface of the premolars and molars displays a pronounced lingual inclination, with the buccal walls being more elevated than the lingual ones. Moderate wear is observed on the teeth on the right maxilla, whereas those on the left display heavy wear. Additionally, root alveoli of the left and right canine and P² can be observed (Figs. 2D, 3D). The anterior part of the premaxilla, including the incisors, is missing. However, an estimated space suggests a relatively broad area for the incisors, confirmed by the reconstruction of the lower jaws by Tafforeau (2004) and Zollikofer et al. (2009).

**Canines** Although not preserved, the large root alveoli are present, measuring approximately 7 mm in mesiodistal length and 6 mm in labiolingual width (Fig. 2D). The canines of *Siamopithecus* have a broadly oval cross-section, with a mesiodistal to labiolingual ratio of approximately 1.16, similar to other primates such as *Bahinia* (1.13; Jaeger et al., 1999), *Pondaungia* (1.21; Shigehara et al., 2002), extant *Tarsius*, platyrrhines, notharctines, and Fayum anthropoids (e.g., *Aegyptopithecus* 1.19; Kay et al., 1981). This configuration differs from adapines and lemuriformes, which typically have more buccolingually compressed canines, except for *Adapis magnus*, which has a ratio of 1.28 (Gingerich and Martin, 1981).

**Premolars** There are three premolars. P² has lost its crown, leaving a small, rounded alveolus positioned more buccally than other premolars. Judging from the size of its alveolus, P² must have been greatly reduced and was probably a peg-like tooth. P³ is smaller compared to P⁴, both displaying triangular outlines in occlusal view. The mesiodistal length on the buccal margin is longer than on the lingual margin. The protocone of P³ is broken, but it is visible on the right tooth (Fig. 4A), and on the holotype TF 3635 (Fig. 4B), in which it is small and low. Both teeth are bicuspid with a well-developed paracone, which is higher and larger than the
protocone. However, P⁴ has a larger protocone than P³. Neither premolar possesses a hypocone, metacone, or conules. The lingual part of the tooth is bordered by the mesial and distal cingula, with a short mesial cingulum present. Neither premolar possesses buccal nor lingual cingulum. P³ and P⁴ have three roots, two buccal and one lingual. The mesiodistal lengths of P³⁻⁴ represent approximately 35% of the P³–M³ length.

**Molars** The molars decrease in size from M¹ to M³. These upper molars are wide buccolingually with well-developed, strongly slanted buccal and lingual walls. M¹ and M² display a trapezoidal occlusal outline, whereas M³ presents a triangular outline. Each molar has four distinct cusps, with the buccal cusps being larger and higher than the lingual cusps. There is no trace of buccal and lingual cingula. The crown height is maximum on M¹, gradually decreasing on M² and M³. The upper molars have three roots, two buccal and one lingual.

M¹ is the largest tooth with prominent buccal cusps (paracone and metacone) that are nearly equal in size, connected by short postparacrista and premetacrista. The parastyle and metastyle are weakly developed with a shallow notch in the buccal wall. A hypoparacrista connects the paracone to the protocone, while the crista obliqua connected the metacone to the protocone, forming a small trigone basin. Although the protocone and hypocone are rather worn, the hypocone appears slightly smaller and lower than the protocone. It is clearly derived from the distal cingulum as on *Ganlea* and its interpretation as a displaced metaconule (Coster et al., 2013) is therefore refuted. The preprotocrista extends from the protocone and connects to the preparacrista at a parastyle, creating a small mesial fovea. Additionally, a short mesial cingulum extends from the parastyle to the mesiolingual wall of the protocone. A distal cingulum connects the hypocone to the metastyle.

M² displays a molar structure similar to M¹, but with a smaller and lower metacone, positioned more lingually compared to the paracone. This result makes an oblique orientation
of the buccal wall. The buccal wall displays a shallow notch. A crista obliqua connects the metacone to the protocone, unlike $M^2$ of the holotype TF 3635, where a crista obliqua connects the metacone to the hypocone (Fig. 4A, B). However, this unique character found in TF 3635 is likely attributed to individual variation. Due to tooth wear, the cusplet initially described as a duplicated protocone (Chaimanee, 2004) cannot be observed as on the holotype TF3635. The enamel on the lingual wall shows a wrinkled appearance, and there is a mesial cingulum and a trace of distal cingulum.

$M^3$ displays a triangular outline and a highly oblique buccal wall, resulting from a well-developed protocone combined with reduced metacone and hypocone. The paracone is large and exhibits a distinct hypoparacrista. Enamel wrinkles are observed on the lingual wall of $M^3$, similar to $M^2$, potentially indicating a diminishing lingual cingulum. Ducrocq (1999) previously noted a fragment of $M^3$ on the holotype TF 3635 as unreduced and considered it a primitive characteristic. However, in this new specimen, $M^3$ is a complete tooth and appears to exhibit a significant reduction.

**Palate and dental arcade** The palate of TF 7625 is incomplete, missing its distal part, and is relatively shallow and short compared to its width (Figs. 2D, 3D). The anterior segment is broken at the level of the anterior palatine foramen. The distal end of the anterior palatine foramen is visible, suggesting a short length, reaching the level of the mesial part of the canine. Due to the displacement and lingual bend of the maxillae, a symmetric µCT image of the right maxilla was used to reconstruct the horizontal palate, revealing a parabolic shape of the dental arcades (Tafforeau, 2004; Zollikofer et al., 2009) (Fig. 5A). Measurements of the distances between the tooth rows were obtained from this µCT-reconstruction. The dental arcade exhibits a minimum width across the buccal points of $P^3$ (~ 30.5 mm) and a maximum width across the buccal points of $M^2$ (~ 42.2 mm). The attachment area for the tendon of the
superficial masseter muscle is well developed and located between M¹ and M². The robust root of the zygomatic process begins at the level of M¹.

*Siamopithecus* displays a unique combination of characteristics, including a notably short muzzle and strongly parabolic tooth rows. Anteriorly, extending to the distal part of P³, the outer walls of the premaxilla/maxilla bones are parallel but gradually diverge and extend laterally, reaching the root of the zygomatic arch.

**Facial morphology** On the left maxilla, notable features include a prominent portion of the zygomatic bone, found attached to the maxilla in the current position. The relatively short muzzle suggests a sub-vertical orientation of the face, with a pronounced frontated orbit (Fig. 5E). The reconstructed lower jaw confirms the short muzzle. The left maxilla reveals a lower margin of the left orbit, the broad root of the zygomatic (15.0 mm), and its ramus. The maxilla contributes to the lower rim of the orbit, a consistent trait among anthropoids. The orbit is positioned anteriorly, slightly above M¹, indicating a nearly vertical orientation based on the alignment of its lateral rim. The lateral edge of the orbit may display a slight forward angle, possibly influenced by postmortem deformation during fossilization, which might have increased the observed frontation angle.

The zygomatic is wide at its base with a thin posterior lamina, extending from its base to its preserved apex. This thin posterior lamina is oriented internally and is broken, preserved for about 1 mm at the base of the zygomatic and up to 3 mm at its medial part. In lingual view, a robust, vertical postorbital bar extends distally, featuring a thin posterior lamina separated by a small groove, indicating incipient or complete postorbital closure (Fig. 2F), a characteristic of anthropoids (Cartmill, 1980; Deleon et al., 2016). Near the mesial end of the maxilla, at the lower orbit rim, a small infraorbital foramen is observable. In lateral view, the dental arcade displays a convex shape, with the highest point located on the buccal wall of M¹. The zygomatic-maxillary suture is identified by a fracture zone extending obliquely from
the base of the orbit to the zygomatic point.

In the frontal view, the left maxillary fragment reveals the anterior part of the maxilla, extending from the mesial anterior extremity of the orbital floor, curving toward the interorbital wall (Fig. 2A). It is broken at a point likely corresponding to the suture with the lacrimal bone, suggesting an internal orbit position for the lacrimal bone, similar to that observed in *Ganlea* (NMMP103). Upon symmetrization, a relatively broad nasal opening is apparent, wider at its inferior level than at its superior level (Tafforeau, 2004) (Fig. 5D).

The right maxilla is less complete compared to the left (Fig. 3). It is incomplete both at its mesial extremity and at its upper range. The orbital region has poor preservation, with only a short segment of its lower lateral rim remaining. The sub-vertical orientation of the zygomatic-maxilla suture line appears oblique, potentially due to a single fracture induced during the fossilization process. The right maxillary fragment preserves the lower part of the zygomatic, which forms the lateral boundary of the orbital cavity. This fragment illustrates the involvement of the maxilla in forming the lower margin of the orbital cavity, a common trait among anthropoids. At the anterior section of the lower orbital margin, there is a break that intersects a small infraorbital foramen, partially preserved in its lower half.

**Orbital floor** The preservation of the left orbital floor is better than that of the right, and the following description is based on its characters (Fig. 2C). The surface displays significant irregularities, making it challenging to accurately assess the impact of post-mortem deformation during fossilization. Primarily, its maxillary bone envelops long premolar and molar roots encased in cancellous bone. This distinctive thickness distinguishes it from strepsirrhines and stands as a defining feature of anthropoids. Furthermore, it is extended distally, forming a substantial surface, a feature absent in strepsirrhines (Shigehara et al., 2002). Towards the lingual aspect and at a lower elevation, an irregular surface emerges,
featuring prominent apices of the molar roots above the surface. Notably, there is an absence
of a parazygomatic groove along the base of the postorbital bar.

**Orbital size** The estimation of orbital size is based on the left maxilla TF 7625, which exhibits better preservation of orbital anatomy compared to the right side. Approximately 18.0 mm of the lower and lateral parts of the orbit rim are preserved on the left maxilla. This length of the preserved orbit rim is sufficient for a reliable estimate of the orbit diameter (Heesy and Ross, 2001). Although the orbit rim is slightly broken in the central part of the preserved portion, the associated missing part and displacement of the rim are minor and thus considered negligible.

An orbit diameter of approximately 15.0 mm was determined for *Siamopithecus*.
Relative to the length of its M¹, the estimated orbit diameter of *Siamopithecus* is
proportionally small compared to tooth size, confidently indicating a diurnal activity pattern
for this anthropoid primate (Fig. 6). This result is consistent with that of Zollikofer et al.
(2009), based on virtual reconstructions of TF 7625. These authors compared the bi-orbital
width of *Siamopithecus* with two skull measurements (C–M³ length, width between the two
M¹s) and inferred a diurnal activity pattern for this anthropoid based on the obtained small
relative orbit size.

**Orbital frontation** According to the virtual reconstruction conducted by Zollikofer et al., (2009), *Siamopithecus* displayed a high frontation angle of 81.6°. It should be noted that this value may be slightly overestimated due to the slight forward orientation of the zygomatic caused by deformation during fossilization. A more accurate estimate might fall within the range of 75°–80°. However, it remains high in any case, indicating a short and sub-vertical facial structure positioned under the orbits.
**Orbital convergence** According to the virtual reconstruction by Zollikofer et al. (2009), *Siamopithecus* displays an average orbital convergence of 72.1°. Although the orbital morphology of *Siamopithecus* is only partially preserved, it suggests a moderate-size orbit relative to its gnathic dimensions, displaying more convergence and frontation compared to other anthropoids. When considering all available data, including orbit diameter, frontation, and convergence values, they indicate a considerably derived stage of crown anthropoid evolution. This interpretation is further supported by the inferred presence of complete or partial postorbital closure in this fossil, as indicated by the remnants of a thin posterior lamina of the zygomatic.

**Mandible** The mandible of *Siamopithecus* (TF 7624) has been previously described by Chaimanee et al. (2000) and Chaimanee (2004), and virtually reconstructed by Tafforeau, (2004), and Zollikofer et al. (2009) (Figs. 5B, C and 7). It is massive and strongly elevated with a nearly vertical (70°) but unfused symphysis (Fig. 7A, B). The height of the symphysis (infradental-gnathion) measures 25.7 mm (Zollikofer et al., 2009). The ramus is missing, being broken 1 cm above the occlusal surface of M<sub>3</sub>. Its anterior part lies behind the distal end of the M<sub>3</sub> talonid, resulting in a short retromolar space equivalent to the length of M<sub>2</sub>.

The canine is robust, with its apex noticeably higher than that of P<sub>3</sub>, and it has an oval-shaped crown outline. It displays a clear wear facet on its distolingual crown made by a strong upper canine. P<sub>2</sub> is broken, showing only a small root, indicating a reduced crown size. P<sub>3</sub> is unicuspidate and mesiodistally longer than wide, unlike those of *Pondaungia* and *Ganlea*, which are nearly as wide as long. Similarly, P<sub>4</sub> is longer than wide, with a small metaconid slightly distal to the protoconid, and its talonid lacks significant development, reduced to a tiny hypoconid cusp. The molars closely resemble those of the paratype (TF 3634) and display more worn and less distinct characters (Fig. 7D, E). They are characterized by the absence of paraconids which are replaced by a preprotocristid that divides an anterior fovea
while connecting to the metaconid. Similar to P₃ and P₄, their buccal walls are slightly slanted and lack a buccal cingulid. All three molars, like those of the paratype, are waisted between trigonid and talonid. The hypoconulid is present on all molars, closer to the hypoconid than to the entoconid, and more strongly developed on the paratype M₁ (Fig. 7D). On M₃, it represents the largest cusps, situated medially. In TF 3634 M₃, which is only slightly worn, numerous small enamel wrinkling crests develop within the basins.

The reconstruction of the maxillae and the facial features (TF 7625) by Tafforeau (2004) also allowed for the reconstruction of the occlusion between the upper and lower dentitions (Fig. 5). This virtual reconstruction provided visualization of the arrangement of the lower dental rows (TF 7624). These rows display a curved shape, widest around the M₂ metaconids, and narrowing towards the area of the canine (Fig. 5B, C).

**Body weight estimation of Siamopithecus** We estimated the body weight of *Siamopithecus* by examining the maxilla of TF 7625. We employed the surface area of M² (mesiodistal length x buccolingual width), due to the variability observed in M¹ within this specimen and applied regression equations provided by Egi et al. (2004). We obtained a predicted body mass for TF 7625 of 7.4 kg. Hence, *Siamopithecus* would have been approximately similar in size to the male white-cheeked gibbon, *Nomascus leucogenys* (Geissmann, 1993).

## 3.2. Comparison

*Siamopithecus* displays several anthropoid characters on its upper molars, such as a single-rooted P², bicuspid premolars, and molars with an almost continuous crista obliqua. Its teeth have a greater buccolingual width compared to mesiodistal length, an unwaisted occlusal outline, a short trigon basin, a hypocone derived from the cingulum, and a well-developed hypoparacrista. In the virtual reconstructions, the upper and lower tooth rows of
*Siamopithecus* present a distinctive parabolic shape unique to this species (Zollikoffer et al., 2009).

**Amphipithecids** *Pondaungia* reveals some similarities in dentition with *Siamopithecus*, such as a connecting crest between the metacone and the hypocone, although this feature is not consistently present in all upper molars of *Siamopithecus* (exceptional on M² of TF 3635). However, there are several distinguishing differences. *Pondaungia* exhibits more pronounced buccolingual development in its premolars, more bunodont molars with less sharp cristae and small vestigial conules (e.g., NMMP 92; Chaimanee et al., 2024), features that are absent in *Siamopithecus*. Additionally, *Pondaungia* has well-developed lingual and buccal cingula on M¹ and M², and M¹ is smaller than M², unlike *Siamopithecus*.

A left maxilla fragment with P³⁻⁴ and M² of *Pondaungia* (NMMP 87; Chaimanee et al., 2024) provides valuable information for comparison with *Siamopithecus*. It exhibits a distinct infraorbital foramen located above P⁴, whereas in *Siamopithecus*, its position is more anteriorly, above P³. The external surface of *Pondaungia*'s maxilla is smooth without a post-canine fossa. However, the tooth row forms a parabolic shape similar to that of *Siamopithecus*. *Pondaungia*'s orbital floor is flat and inclined medially. The lingual roots of M¹ and M² are salient on the orbital floor but to a lesser degree than those of *Siamopithecus*. In internal view, NMMP 87 displays a notably large, deep, and concave depression, indicating a massive maxillary sinus. This sinus is not observable in *Siamopithecus*. Other maxillary fragments of *Pondaungia* (NMMP 12 and NMMP 18) also display a significant maxillary sinus (Takai and Shigehara, 2004). However, unlike the two other *Pondaungia* maxillae, there is no parazygomatic groove present in NMMP 87.

*Ganlea* exhibits more primitive dental features than *Pondaungia* (Beard et al., 2009; Jaeger et al., 2020). It has a smaller M¹ compared to M², a prominent hypocone derived from the distolingual cingulum, a small paraconule, and a complete lingual cingulum on its molars.
Its crista obliqua joins the protocone more distally than in *Siamopithecus*. However, unlike in one M² of *Siamopithecus* and most upper molars of *Pondaungia*, it does not connect to the hypocone. *Myanmarmpithecus* (Takai et al., 2001) also has molars displaying the same organization as those of *Ganlea*.

A cranial fragment of *Ganlea* (NMMP103; Jaeger et al., 2020) provides valuable insights for comparison with *Siamopithecus*. The *Ganlea* palate shows divergent tooth rows. It also exhibits a high estimated orbit convergence (72.2°), similar to that of *Siamopithecus* (72.1°). The orbit size of *Ganlea* is estimated to be 15.56 mm in diameter (Jaeger et al., 2020) and 15.0 mm in *Siamopithecus*. *Ganlea* has an elongated rostrum that accommodates its very large upper canines, contrasting with *Siamopithecus*. The external surface of the *Ganlea* maxilla is not smooth and displays a post-canine fossa, which may be related to the enlarged canines in *Ganlea*. The frontal bone in *Ganlea* indicates an absence of postorbital closure. The section of the base of its postorbital bar is triangular and mesiodistally short, unlike the flattened and extended posterior lamina seen in *Siamopithecus*, which suggests a partial or complete postorbital closure in the latter. *Ganlea* has an orbital floor that differs noticeably, being flat, shorter and inclined medially, similar to those of *Pondaungia*. Unlike *Siamopithecus*, there is no observable prominent root tip on the *Ganlea* orbital floor.

The similarities between *Ganlea* (NMMP 103) and *Pondaungia* (NMMP 87) align with their close phylogenetic relationships, reflecting common anatomical features among Myanmar Eocene anthropoids. These include short faces and relatively thick maxillae, characters also shared by other Asian anthropoids like *Bahinia*. However, *Siamopithecus* exhibits distinct anatomical features not observed in other known Asian anthropoids. For instance, its molars have root tips protruding noticeably on the orbital floor (Fig. 2C), a feature less prominent in *Ganlea* and *Pondaungia*. Additionally, *Siamopithecus* displays a thin bony lamina on the distal part of its zygomatic bone, indicating the potential presence of
a partial or complete postorbital wall, which is not observed in other Asian Eocene anthropoids. There is no indication of postorbital closure in *Pondaungia*. However, given its close phylogenetic relationships with *Ganlea*, an optimized inference would support a lack of closure as in *Ganlea*.

*Aseanpithecus* displays several differences. These include a more extensive buccolingual extension of premolars, the presence of both buccal and lingual cingula surrounding the molars, less inclined molar walls, broader and larger trigons, and lack of hypocone. *Aseanpithecus* has been identified as an Asian anthropoid that shares a striking resemblance in its molar anatomy with African oligopithecids (Jaeger et al., 2019).

**Propliopithecids** The upper molars of Taqah propliopithecid (TQ3), from the early Oligocene of Taqah, Oman, initially described as *Moeripithecus markgrafi* by Thomas et al. (1991), exhibit both similarities and distinctions compared to those of *Siamopithecus*. Significant resemblances include the larger size of M¹ compared to M², reduced bunodonty, conule absence, strong crista obliqua, and weak hypocones. Differences include *Siamopithecus* lacking a prominent lingual cingulum and having more slanted cups. TQ3 displays less developed and elevated hypocones, lacks connection crests between the protocone and hypocone, and has a strong lingual cingulum with a pericone on its molars, contrasting with *Siamopithecus*. Additional differences are observed in premolar morphology. TQ3 lacks P² and displays an enlarged P³, with greater buccolingual extension, with robust protocones. Both P³ and P⁴ exhibit transverse crests connecting the protocone to the paracone, forming anterior, median, and posterior fovea, a feature not found in *Siamopithecus*. Furthermore, TQ3 develops lingual cingula, which are absent in *Siamopithecus*. Notably, P³ is larger than P⁴ in TQ3, contrary to *Siamopithecus*, suggesting a potentially more primitive premolar development for *Siamopithecus*.
The lower tooth row shares many morphological characters with those of the Taqah propliopithecid (TQ4). The molar structure is similar, with differences including the less central position of the hypoconulids, their smaller size on M<sub>2</sub>, the absence of buccal cingulids, the less bunodont cuspids, and the larger development of the M<sub>3</sub> hypoconulid in *Siamopithecus*. However, as with the upper dentition, the premolars exhibit more significant differences. The P<sub>3</sub> of TQ4 is notably larger than P<sub>4</sub>, and its P<sub>4</sub> has a well-defined talonid basin and a stronger metaconid, positioned at the same level as the protoconid rather than distal to it as in *Siamopithecus*. This character has been cited as a synapomorphy of crown anthropoids and is lacking in *Siamopithecus*.

Taqah propliopithecid (TQ3 and TQ4) is generally recognized as the basal sister group of the more derived Fayum propliopithecids (Seiffert, 2012), but its peculiar characters, such as the small size of its molar hypocones, render its phylogenetic position uncertain. However, the resemblance of its dentition to that of *Siamopithecus* raises interesting questions relative to their phylogenetic relationships. *Siamopithecus* appears by some of its characters to be more primitive (premolar structure and retention of P2) and by others more specialized (inclined molar walls and crest linking protocone and hypocone).

*Aegyptopithecus* is a highly derived anthropoid with a closed postorbital wall. *Siamopithecus* differs from it in several significant aspects. The tooth rows of *Siamopithecus* form a parabolic shape rather than diverging posteriorly, as in *Aegyptopithecus* (Simons, 1987; Leakey et al., 1991). *Siamopithecus* displays a shorter muzzle compared to that of *Aegyptopithecus* (Leakey et al., 1991). *Siamopithecus* exhibits higher frontation (81.2° in *Siamopithecus*; Zollikofer et al., 2009; 69° in *Aegyptopithecus*; Simons, 2004) but a lesser convergence angle (72.1° x 2 in *Siamopithecus*; Zollikofer et al., 2009; 183–192° in *Aegyptopithecus*; Simons, 2008) and displays a smaller orbit (15.0 mm) compared to *Aegyptopithecus* (16.9–18.9 mm; Kay and Kirk, 2000). Another distinguished feature is the
anterior location of the contact point between the zygomatic and maxillary bone on the lower
orbit rim in *Siamopithecus*, indicating more anteriorly positioned orbits compared to
*Aegyptopithecus*. Furthermore, the zygomatic branch of *Siamopithecus* begins at the level of
*M¹*, whereas in *Aegyptopithecus*, the orbit is located more posteriorly above *M²*, and the
elevation of the maxilla is higher (Simons, 1987). The ethmoid region is not preserved in
*Siamopithecus* and therefore cannot be compared to that of *Aegyptopithecus* (Rossie et al.,
2002; Rossie, 2005).

**Pliopithecids** Few pliopithecid crania are preserved. Among them, *Epipliopithecus vindobonensis* (Individual II) represents a derived Middle Miocene representative of the pliopithecids (Zapfe, 1960). This specimen shows similarities with *Siamopithecus*, such as orbit size and anterior position, a vertically oriented face, high frontation and convergence values, a short muzzle, a wide base of the zygomatic root, and comparable orbital floor organization. Despite these similarities, the presence of $P^2/2$ and the absence of shared derived dental characters does not allow us to consider *Siamopithecus* as a pliopithecid.

**Parapithecids** The two parapithecid crania from Fayum deposits, *Parapithecus* (DPC 18651) and *Apidium* (DPC 9867), exhibit complete postorbital closure and possess numerous anatomical differences that distinguish them from *Siamopithecus*. For instance, their facial structure is not as short, and their frontation and convergence angles are notably lower. The convergence angle of *Parapithecus* (99-106°) and *Apidium* (105-113°) (Simons, 2008) is less than that of *Siamopithecus* (72.1° x 2 by Zollikofer et al., 2009). Similarly, the frontation angle of *Parapithecus* is 62° (Simons, 2004), lower than that observed in *Siamopithecus* (81.6°; Zollikofer et al., 2009). Additionally, both parapithecids display a prominent zygomasseteric foramen, a characteristic not observed in *Siamopithecus*. In occlusal view, their dental arcade is less parabolic, and the palate length/breath index is considerably larger compared to *Siamopithecus*.
In summary, *Siamopithecus* displays a combination of primitive and derived characters not commonly observed in other anthropoids. Primitive traits include the absence of conules, the presence of P²/₂, and the small sizes of P³ and its protocone. Conversely, more derived features include the inclined orientation of the buccal and lingual walls on the upper molars, the presence of a crest formed by a portion of the postprotocrista and the prehypocrista connecting the two lingual cusps, and the absence of both buccal and lingual cingula on premolars and molars. These characters, associated with their unique facial and orbital morphology, highlight its distinctiveness among anthropoids.

### 3.3. Phylogenetic affinities

Numerous phylogenetic analyses have suggested that *Siamopithecus* is an amphipithecid, either as a sister group to the Myanmar amphipithecid *Pondaungia*, *Myanmarpithecus*, and *Ganlea* or phylogenetically close to this clade (Kay et al., 2004; Seiffert et al. 2005, 2018; Beard et al. 2009; Chaimanee et al., 2012, 2013; Marivaux et al. 2013, 2023; Jaeger et al. 2019). Based on their phylogenetic results and the greater shared morphological features within the Myanmar amphipithecid, Beard et al. (2009) proposed classifying *Siamopithecus* into the subfamily Siamopithecinae and the three Myanmar amphipithecid (*Pondaungia*, *Myanmarpithecus*, and *Ganlea*) into the subfamily Amphipithecinae. However, with the recovery of more complete remains of the amphipithecine *Ganlea* (Jaeger et al., 2020), closer comparisons have revealed numerous differences between *Ganlea* and *Siamopithecus*. The main differences concern molar structure, premolar organization, snout length, and the presence of a postorbital closure. Since no postorbital closure is present in *Ganlea*, the hypothesis of a postorbital reopening among amphipithecines seems very unlikely, as such an evolutionary reversal has never been documented among primates.
Therefore, we reinvestigated the phylogenetic position of *Siamopithecus* using maximum parsimony analysis with PAUP 4.3.99.169.0 (Swofford, 2003) with three different data matrices. In our first analysis, we employed a matrix including 117 taxa and 394 morphological characters based on the data matrix of Seiffert et al. (2018). Conducting a heuristic search in PAUP using the original settings of Seiffert et al. (2018) (utilizing a backbone tree derived from the molecular phylogeny of extant strepsirrhines, weighted characters, and 4 characters with irreversible states), we obtained 20 most parsimonious trees (length = 4727.5, CI = 0.1518, RI = 0.5590). In the strict consensus tree (Fig. 8), *Siamopithecus* is placed within the amphipithecids and is a sister group to the amphipithecines. The amphipithecids are positioned as a sister group to the crown-anthropoids.

In the second analysis, we used the data matrix from Jaeger et al. (2020), enhanced with the new characters for *Siamopithecus*. This data matrix includes 38 taxa and 326 morphological characters (both dental and osteological). Five most parsimonious trees (length = 1425 steps, CI = 0.3411, RI = 0.5581) were obtained after a heuristic search (random addition of taxa, 1000 replications, unweighted characters, no topological constraint). *Siamopithecus* is placed within the amphipithecid clade (see Supplemental Online Material; Fig. S1). In the majority-rule consensus tree, the amphipithecids form a clade of stem anthropoids, similar to the results of Jaeger et al. (2020).

In the third analysis, the data matrix, derived from those of Jaeger et al. (2019, 2020), included a broader sample of fossil anthropoids (45 taxa in total) and a similar set of characters. Twelve most parsimonious trees (length = 1514 steps, CI = 0.3210, RI = 0.5554) were obtained after a heuristic search with the same parameters as in the first analysis. In the strict consensus tree (see Supplemental Online Material; Fig. S2), *Siamopithecus* is part of a poorly resolved clade that included catarrhines, platyrrhines, parapithecoids, oligopithecids
and Asian anthropoids commonly classified as amphipithecids (*Pondaungia*, *Ganlea*, *Myanmarpithecus*, *Krabia*). The amphipithecids are not monophyletic in this tree. However, in the 50% majority consensus tree, *Siamopithecus* is again grouped with the amphipithecids clade together with *Pondaungia*, *Ganlea*, *Myanmarpithecus*, and *Krabia*, and amphipithecids are reconstructed as crown anthropoids.

These results are generally inconsistent with our morphological analysis of the skull and dentition of *Siamopithecus*, which suggests that amphipithecids are likely not monophyletic. All of these results also indicate a homoplastic evolution of the postorbital closure in anthropoids, presenting an unparsimonious scenario for this key morphological feature. Additionally, we observe that the phylogenetic position of the amphipithecids varies in our results, similar to previous phylogenetic analyses (see e.g., Seiffert et al. 2005, Beard et al. 2009, Jaeger et al. 2020). The results from the third analysis contradict those of Jaeger et al. (2020), which suggested that the amphipithecids were stem anthropoids with a more basal position than the advanced stem anthropoids (such as parapithecoids). The results from the first analysis are also improbable, as they imply a loss of the postorbital closure in the amphipithecines.

We believe that the large number of derived (and possibly homoplastic) dental traits observed in amphipithecids compared to cranial/postcranial traits creates a bias in the analysis. Aside from *Aseanpithecus*, only *Ganlea* and *Siamopithecus* are the Paleogene Asian anthropoids with available non-dental-gnathic features, and *Ganlea* is the only taxon with firmly associated postcranial material. Although important, the new cranial features documented for *Siamopithecus* constitute a small fraction of the total number of characters for this anthropoid (less than 2% in the first analysis, and less than 3% in the second and third analyses). As a result, the retained topologies for anthropoids remain inadequately constrained
by the fossil record. Therefore, we believe that our parsimony analyses do not provide reliable
affinities for *Siamopithecus*.

**4. Discussion and conclusion**

The reexamination of *Siamopithecus* maxilla and facial fragment confirms its
classification as an anthropoid, supported by numerous cranial and dental characters
previously documented by Zollikofer et al. (2009). Newly identified characters, particularly
the potential presence of partial or complete postorbital closure, necessitate a reassessment of
its phylogenetic affinities. Although its status as an anthropoid is clear, in opposition to
Rosenberger et al. (2011) interpretation, its precise phylogenetic position within the
anthropoids remains unsolved. Comparative analysis with recently discovered cranial
fragments of amphipithecines, particularly *Ganlea* (NMMP103) and *Pondaungia* (NMMP87),
reveals notable distinctions despite some shared dental features. These similarities include the
connection between the protocone and hypocone on upper molars, weakly developed or
absent conules, enamel wrinkling, and thin enamel without Hunter-Schreger bands. Both
*Ganlea* and *Pondaungia* also exhibit an elevated mandibular corpus, indicative of a hard-food
diet (Smith, 1983; Ramdashan et al., 2010).

The presence of partial or complete postorbital closure in *Siamopithecus* excluded it
from being classified among the amphipithecines. The absence of postorbital closure is a
distinguishing feature of amphipithecines, known from *Ganlea* and inferred for other
amphipithecines through phylogenetic optimization (Chaimanee et al., 2024). *Siamopithecus*
also lacks distinctive shared derived characters typical of amphipithecines, such as the
pronounced buccolingual extension of the lower premolars.

In comparison to eosimiids (including *Bahinia*), *Siamopithecus* premolars and molars
display more derived characters, including a well-developed cingular-derived hypocone in the
upper molars, the absence of paraconids in the lower molars, and the presence of a metaconid on P4. Therefore, the establishment of a new family, *Siamopithecidae*, is justified. In this study, *Siamopithecus* is not considered a basal sister group of amphipithecines due to the probable presence of postorbital closure, challenging the hypothesis that amphipithecines lost their postorbital closure. Such a hypothesis lacks parsimony and is unsupported by evidence from primates. Consequently, shared dental morphology and dietary adaptations between *Siamopithecus* and amphipithecines are interpreted as homoplasies rather than shared derived characters.

Phylogenetic analyses over the past two decades have proposed two main positions for *Siamopithecus*. Initial studies, primarily based on dental characters, placed it within the amphipithecids, often as the basal sister clade of amphipithecines. Amphipithecids were interpreted as members of the crown anthropoids, either as a sister group to catarrhines (Beard et al. 2009; Chaimanee et al., 2012; Jaeger et al., 2019) or sometimes to platyrhines (Seiffert et al., 2010; Marivaux et al. 2023). However, subsequent discoveries of more complete amphipithecines remains (Jaeger et al., 2020) strengthen support for placing amphipithecines as stem anthropoids (Marivaux et al., 2005; Seiffert et al., 2005; Marivaux, 2006; Ni et al., 2013; Jaeger et al., 2020), with *Siamopithecus* sometimes resolved as their closest outgroup (Seiffert et al., 2005, 2018; Seiffert, 2012; Jaeger et al., 2020). One exceptional result placed *Siamopithecus* as a sister group of simiiforms, outside the Amphipithecinae, assigned to a more basal clade (Boyer et al., 2010; Coster et al., 2013).

New data suggest alternative potential phylogenetic relationships for *Siamopithecus*. It shares several characters with the most primitive known Taqah propliopithecid, described by Thomas et al. (1991). In contrast, Taqah propliopithecid has significantly more derived premolars, with P³/₃ larger than P⁴/₄ and a more robust protocone on P³. The primitive structure of *Siamopithecus* premolars and the persistence of small, reduced P²/₂ suggest that it
has not attained the catarrhine status. Therefore, a hypothesis of a stem catarrhine
phylogenetic position for *Siamopithecus* can be proposed. The origin of propliopithecid
remains controversial. Most authors suggest that propliopithecid derived from African
Eocene anthropoids in an undocumented Afro-Arabian province (Simons, 1990; Seiffert et al.,
2004). The absence of African ancestors of propliopithecid in the rich late Eocene Fayum
faunas (Seiffert, 2012) and their sudden appearance during the early Oligocene in Afro-Arabia
remains an enigma. In Taqah (Oman), the early Oligocene propliopithecid is associated with
cricetid rodents (Thomas et al., 1999; Sen, 2012), which are undoubtedly of Asian origin
(Rodrigues et al., 2010). This association raises the hypothesis that propliopithecid may have
originated in Asia, similar to cricetid rodents. In this context, *Siamopithecus* may document a
primitive relative of these unknown Asian propliopithecid. The discovery of Eocene
propliopithecid in Afro-Arabia would prompt a reconsideration of this interpretation and
encourage the search for new phylogenetic relationships between *Siamopithecus* and African
Paleogene anthropoids.

In conclusion, recent discoveries related to Myanmar amphipithecines have allowed more
precise comparisons of their maxillae, revealing significant anatomical differences between
amphipithecines and *Siamopithecus*. These findings have led to the reinterpretation of shared
derived dental and mandibular characters associated with a hard-food diet as homoplasies.
Consequently, *Siamopithecus* has been reclassified outside of the amphipithecid and
attributed to a new family, the Siamopithecidae. Based on dental similarities between
*Siamopithecus* and the most primitive known propliopithecid, from Oman, a phylogenetic
position within the stem catarrhines is proposed. This would implicate an Asian origin for
propliopithecid rather than deriving from oligopithecid ancestors in an undocumented Afro-
Arabian province. However, the status of the Taqah propliopithecid from Oman remains
uncertain, and further discoveries should validate or challenge this revised interpretation.
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## Acknowledgments

The authors would like to thank Christoph Zollikofer (University of Zurich) for providing the comparative materials. We extend our gratitude to Paul Tafforeau and European Synchrotron Radiation Facility (ESRF) beamlines ID 17 for the CT-scan pictures (Fig. 4, 5, and 7), and ID 19 (Fig. 4B), Sabine Riffaut (PALEVOPRIM) for creating the figures, and the Electricity Generating Authority of Thailand (EGAT) for their scientific assistance and access to the mine. We also thank Mark Grabowski (the editor), the associated editor, and two anonymous reviewers, Laurent Marivaux (ISEM) and K. Christopher Beard (University of Kansas) for their helpful comments and suggestions. This work has been supported by the CNRS-UMR 7262, the University of Poitiers, and the Department of Mineral Resources, Thailand (DMR). Partially funding was provided by the ANR-18-CE92-0029 Project (to O. Chavasseau).
<table><caption>**Table 1** Dental measurements of *Siamopithecus eocaenus* (MD, mesiodistal length; BL, buccolingual width; trig, trigonid; tal, talonid). Maxilla (TF 7625) compared with TF 3635* (holotype, Chaimanee et al., 1997) and mandible (TF 7624#, Chaimanee et al., 2000, and TF 3634+, Chaimanee et al., 1997) (in mm).</caption><thead><tr><th></th><th colspan="2">TF 7625<br>Left (worn)</th><th colspan="2">TF 7625<br>Right</th><th colspan="2">TF 3635*<br>Holotype</th><th colspan="2">TF 7624#</th><th colspan="2">TF 3634+</th></tr><tr><th></th><th>MD</th><th>BL</th><th>MD</th><th>BL</th><th>MD</th><th>BL</th><th>MD</th><th>BL<br>(trig/tal)</th><th>MD</th><th>BL<br>(trig/tal)</th></tr></thead><tbody><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>C</td><td>6.46</td><td>4.96</td><td>-</td><td>-</td></tr><tr><td>P<sup>3</sup></td><td>4.4</td><td>5.3</td><td>4.8</td><td>5.8</td><td>4.3</td><td>5.3</td><td>P<sub>3</sub></td><td>4.78</td><td>5.46</td><td>-</td><td>-</td></tr><tr><td>P<sup>4</sup></td><td>4.8</td><td>7.0</td><td>5.0</td><td>7.0</td><td>5.0</td><td>6.5</td><td>P<sub>4</sub></td><td>5.36</td><td>5.77</td><td>-</td><td>-</td></tr><tr><td>M<sup>1</sup></td><td>5.7</td><td>8.6</td><td>5.8</td><td>9.0</td><td>5.4</td><td>8.1</td><td>M<sub>1</sub></td><td>6.37</td><td>6.12/5.96</td><td>-</td><td>-/5.7</td></tr><tr><td>M<sup>2</sup></td><td>5.0</td><td>8.9</td><td>5.1</td><td>9.2</td><td>4.8</td><td>8.1</td><td>M<sub>2</sub></td><td>6.04</td><td>5.92/5.32</td><td>6.5</td><td>5.7</td></tr><tr><td>M<sup>3</sup></td><td>4.5</td><td>7.1</td><td>4.6</td><td>7.5</td><td>-</td><td>-</td><td>M<sub>3</sub></td><td>7.97</td><td>5.22/4.22</td><td>8.1</td><td>5.1</td></tr><tr><td>P<sup>3</sup>-M<sup>3</sup></td><td>22.8</td><td></td><td>25.0</td><td></td><td>-</td><td></td><td>P<sub>3</sub>-M<sub>3</sub></td><td></td><td>29.69</td><td></td><td>-</td></tr><tr><td>M<sup>1</sup>-M<sup>3</sup></td><td>14.7</td><td></td><td>15.7</td><td></td><td>-</td><td></td><td>M<sub>1</sub>-M<sub>3</sub></td><td></td><td>19.32</td><td></td><td>-</td></tr><tr><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>C-M<sub>3</sub></td><td></td><td>40.0</td><td></td><td>-</td></tr></tbody></table>
<table><caption>**Table 2** Measurements of the cranium of *Siamopithecus eocaenus* (TF 7625).</caption><tr><td>Estimated orbit frontation*</td><td>81.6°</td></tr><tr><td>Estimated orbit convergence*</td><td>72.1°</td></tr><tr><td>Biorbital width (Bi-frontomolare orbitale)*</td><td>52.9 mm</td></tr><tr><td>Orbit diameter</td><td>15.0 mm</td></tr><tr><td>Palate width at M<sup>2</sup> (between lingual alveolar borders of M<sup>2</sup>)*</td><td>22.9 mm</td></tr><tr><td>Breadth across P<sup>3</sup> (between buccal alveolar borders of P<sup>3</sup>)*</td><td>30.5 mm</td></tr><tr><td>Breadth across M<sup>2</sup> (between buccal alveolar borders of M<sup>2</sup>)*</td><td>42.2 mm</td></tr><tr><td>Breadth across M<sup>3</sup> (between buccal alveolar borders of M<sup>3</sup>)*</td><td>40.6 mm</td></tr><tr><td>Elevation of maxillary bone on vertical section</td><td>14.5 mm</td></tr></table>
<figure><img src="image_3.png" /></figure>

**Fig. 1** Map of the Krabi coal mine (Krabi basin, Peninsular Thailand), showing the three coal pits (Bang Pu Dum, Bang Mark, and Wai Lek). Fossils of *Siamopithecus* TF 3635 (maxilla, holotype), TF 7624 (mandible), and TF 7625 (maxilla in this study) were recovered from the Bang Mark pit, while TF 3634 (partial mandible) was recovered from the Wai Lek pit.
<figure><img src="image_4.png" /><figcaption>**Fig. 2** The left maxilla of *Siamopithecus eocaenus* (TF 7625), rendered from a 3D virtual model generated from µCT scans, in mesial (A), jugal (B), superior (C), occlusal (D), distal (E), and lingual (F) views. Thin bony lamina on the distal part of its zygomatic, indicating the potential presence of a partial or complete postorbital wall (red arrows).</figcaption></figure>
<figure><img src="image_5.png" /><figcaption>**Fig. 3** The right maxilla of *Siamopithecus eocaenus* (TF 7625), rendered from a 3D virtual model generated from µCT scans, in mesial (A), jugal (B), superior (C), occlusal (D), distal (E), and lingual (F) views.</figcaption></figure>
<figure><img src="image_6.png" /></figure>

<figure><img src="image_7.png" /><figcaption>**Fig. 4** Comparison of right maxillae of *Siamopithecus eocaenus* (TF 7625) (A) and the TF 3635 (holotype) (B), rendered from a 3D virtual model generated from µCT scans (Tafforeau, 2004; https://paleoanthro.org/static/dissertations/Paul%20Tafforeau-lowres.pdf) with the author's authorization). In TF 3635, M² differs by having a transversely oriented hypometacrista that extends lingually from the metacone toward the hypocone (red arrow).</figcaption></figure>
<figure><img src="image_8.png" /><figcaption>**Fig. 5** Virtual reconstitution of the maxilla of *Siamopithecus eocaenus* (TF 7625) using the symmetry of the right maxilla on the horizontal palate in occlusal view (A). The mandible of *Siamopithecus eocaenus* (TF 7624) is reconstructed through the symmetry of the right mandible in the inferior (B) and occlusal (C) views. A virtual reconstitution of the face of *Siamopithecus eocaenus* is achieved by mirroring the left maxilla of TF 7625 and the right mandible of TF 7624, anterior (D), and right lateral (E) views. Images are digital renderings of 3D virtual models generated from µCT scan data from Tafforeau, 2004; https://paleoanthro.org/static/dissertations/Paul%20Tafforeau-lowres.pdf) with the author's authorization).</figcaption></figure>
<figure><img src="image_9.png" /><figcaption>**Fig. 6** Bivariate plot of log-transformed orbit diameter versus first molar length in a sample of extant and fossil primates. *Siamopithecus* displays a small orbit diameter relative to molar size, indicating a diurnal activity pattern. Orbit diameters for extant primates are mediolateral diameters. Data sources from Heesy and Ross (2001) and Jaeger et al. (2019, 2020).</figcaption></figure>
<figure><img src="image_10.png" /><figcaption>**Fig. 7** *Siamopithecus eocaenus* rendered from a 3D virtual model generated from µCT scans (Tafforeau, 2004; https://paleoanthro.org/static/dissertations/Paul%20Tafforeau-lowres.pdf) with the author's authorization, right mandible with C-M₃ (TF 7624) in buccal (A), lingual (B), inferior (C), and occlusal (D) views and right mandible with distal M₁-M₃ (TF 3634) in occlusal view (E).</figcaption></figure>
<figure><img src="image_11.png" /><figcaption>**Fig. 8** Phylogenetic position of *Siamopithecus* among the primates. Results of the maximum parsimony analysis based on a taxon-character data matrix of 394 characters and 117 taxa based on the data matrix of Seiffert et al. (2018) (SOM). A Strict consensus tree of 20 most parsimonious trees (length = 4727.5 steps, CI = 0.1518, RI = 0.5590). The numbers above the nodes are the frequencies of the clade among the most parsimonious trees.</figcaption></figure>
**Supplementary Online Material (SOM):**

Maxilla of *Siamopithecus eocaenus* (Anthropoidea, Primates) from the Paleogene of Krabi, Thailand, and its taxonomic status

Yaowalak Chaimanee<sup>a,*</sup>, Sasa-On Khansubha<sup>b</sup>, Olivier Chavasseau<sup>a</sup>, Arnaud Mazurier<sup>c</sup>, Jean-Jacques Jaeger<sup>a</sup>

<footnote><sup>a</sup>Laboratory Paleontology, Evolution, Paleoenvironments and Paleoprimatology (PALEVOPRIM), UMR 7262 CNRS, University of Poitiers, France.</footnote>

<footnote><sup>b</sup>Department of Mineral Resources, Rama VI Road, Bangkok 10400, Thailand.</footnote>

<footnote><sup>c</sup>Institute of Chemistry of Environments and Materials of Poitiers (IC2MP), UMR 7285 CNRS, University of Poitiers, France.</footnote>

<footnote>*Corresponding author*: E-mail address: Y. Chaimanee [yao.chaimanee@univ-poitiers.fr](mailto:yao.chaimanee@univ-poitiers.fr)</footnote>
# Phylogenetic analyses

## Analysis 1

In this analysis, the data matrix is based on Seiffert et al. (2018), augmented with new coding for *Siamopithecus* (7 new characters). The data matrix includes 117 taxa (33 fossil and living anthropoids, 3 living and fossil tarsiids, 19 omomyids, and 55 fossil and extant strepsirrhines) and 394 morphological characters (see character-taxon-matrix Analysis 1).

A heuristic search was performed in PAUP 4.0a169 using the hsearch command with a random addition sequence of taxa (100 replications), 178 characters were assigned a weight of 1, 216 characters at 0.5, 261 characters were treated as ordered and 4 characters as irreversible. The retrieved topologies were constrained by a backbone tree based on the molecular phylogeny of extant strepsirrhines. All search options follow the protocol used by Seiffert et al. (2018) (Fig. 8).

## Analysis 2

In this analysis, the data matrix is based on Jaeger et al. (2020) with the coding of *Siamopithecus* augmented by newly available features (9 new characters states). The data matrix includes 38 primate taxa (27 anthropoids, 2 tarsiids, 4 omomyids, and 5 adapiform strepsirrhines) and 326 cranial, dental, and postcranial characters (see character-taxon-matrix Analysis 2).

A heuristic search was performed in PAUP 4.0a169 using the hsearch command with a random addition sequence of taxa (1000 replications). No character weighting was used, and 144 characters were treated as ordered (Fig. S1).
<figure><img src="image_12.png" /><figcaption>**Fig. S1** Majority-rule consensus tree derived from the 5 most parsimonious trees obtained in Analysis 2 (length = 1425 steps, CI = 0.3411, RI = 0.5581). The clade frequencies among the most parsimonious trees are indicated.</figcaption></figure>
Analysis 3

In this analysis, the data matrix is based on those of Jaeger et al. (2019, 2020), incorporating the new coding of *Ganlea* from Jaeger et al. (2020). The data matrix includes 45 primate taxa (34 anthropoids, 2 tarsiids, 4 omomyids, and 5 adapiform strepsirrhines) and 326 cranial, dental, and postcranial characters (see character-taxon-matrix Analysis 3).

A heuristic search was performed in PAUP 4.0a169 with the hsearch command with a random addition sequence of taxa (1000 replications). No character weighting was used and 144 characters were treated as ordered (Fig. S2).
<figure><img src="image_13.png" /><figcaption>**Fig. S2** Majority-rule consensus tree derived from the 12 most parsimonious trees obtained in Analysis 3 (length = 1514 steps, CI = 0.3210, RI = 0.5554). The clade frequencies among the most parsimonious trees are indicated.</figcaption></figure>
Character-taxon-matrix

Analysis 1: We use a matrix with 117 taxa and 394 morphological characters based on
Seiffert et al. (2018).

#NEXUS
BEGIN TAXA;
DIMENSIONS NTAX=117;
TAXLABELS
'Tupaia glis'
'Absarokius sp.'
'Abuqatrania basiodontos'
'Adapis parisiensis'
'Aegyptopithecus zeuxis'
'Afradapis longicristatus'
'Aframonius dieides'
'Algeripithecus minutus'
'Allenopithecus nigroviridis'
'Alouatta seniculus'
'Altanius orlovi'
'Anchomomys frontanyensis'
'Anchomomys gaillardi'
'Anchomomys milleri'
'Aotus trivirgatus'
'Apidium moustafai'
'Apidium phiomense'
'Arapahovius gazini'
'Artocebus calabarensis'
'Arsinoea kallimos'
'Asiadapis cambayensis'
'Azibius trerki'
'Bahinia pondaungensis'
'Bahinia banyueae'
'Biretia fayumensis'
'Biretia megalopsis'
'Branisella boliviana'
'Bugtilemur mathesoni'
'Caenopithecus lemuroides'
'Cantius abditus'
'Cantius ralstoni'
'Carpolestes simpsoni'
'Catopithecus browni'
'Cheirogaleus major'
'Darwinius masillae'
'Djebelemur martinezi'
'Dolichocebus gaimanensis'
'Donrussellia provincialis'
'Dyseolemur pacificus'
'Eosimias centennicus'
'Europolemur dunaifi'
'Europolemur klatti'
'Galagoides demidoff'
'Galago moholi'
'Ganlea megacanina'
'Gatanthropus micros'
'Hemiacodon gracilis'
'Hoanghonius stehlini'
'Karanisia clarki'
'Komba spp.'
'Laomaki yunnanensis'
'Lemur catta'
'Leontopithecus rosalia'
'Lepilemur mustelinus'
'Leptadapis magnus'
'Leptadapis priscus'
'Leptadapis ruetimeyeri'
'Loris tardigradus'
'Loveina zephryi'
'Macrotarsius montanus'
'Mahgarita stevensi'
'Marcgodinotius indicus'
'Masradapis tahai'
'Mazateronodon endemicus'
'Mescalerolemur horneri'
'Microadapis sciureus'
'Microcebus murinus'
'Microchoerus erinaceus'
'Muangthanhinius siami'
'Myanmarpithecus yarshensis'
'Nannopithex abderhaldeni'
'Nannopithex raabi'
'Necrolemur spp.'
'Nievesia sossisensis'
'Nyxticeboides simpsoni'
'Nyxticebus coucang'
'Oligopithecus rogeri'
'Oligopithecus savagei'
'Omomys sp.'
'Otolemur crassicaudatus'
'Pan troglodytes'
'Parapithecus fraasi'
'Parapithecus grangeri'
'Perodicticus potto'
'Phenacopithecus xueshii'
'Plesiadapis tricuspidens'
'Plesiolestes problematicus'
'Plesiopithecus teras'
'Pondaungia cotteri'
'Pronycticebus gaudryi'
'Propithecus spp.'
'Propliopithecus chirobates'
'Proteopithecus sylviae'
'Protoadapis curvicuspidens'
'Pseudoloris parvulus'
'Ptilocercus kylin'
'Purgatorius unio'
'Qatrania wingi'
'Rencunius zhoui'
'Saharagalago misrensis'
'Saimiri sciureus'
'Serapia eocaena'
'Shoshonius cooperi'
'Siamopithecus eocaenus'
'Steinius spp.'
'Taqah propliopithecid'
'Tarsius bancanus'
'Tarsius sirindhornae'
'Teilhardina americana'
'Teilhardina asiatica'
'Teilhardina belgica'
'Tetonius sp.'
'Varecia variegata'
'Wadilemur elegans'
'Washakius insignis'
'Xanthorhysis tabrumi'
'Yunnanadapis folivorus'
;
ENDBLOCK;

BEGIN CHARACTERS;
DIMENSIONS NCHAR=394;
FORMAT DATATYPE = STANDARD GAP = - MISSING = ? SYMBOLS = "012345678";
CHARLABELS
[1] 'lower incisor number (modified from Ross et al. 1998 character i1)'
[2] 'lower incisor occlusal arrangement (Ross et al. 1998 character i2)'
[3] 'toothcomb (modified from Ross et al. 1998 character i13)'
[4] 'lower i1-2 area relative to lower m1 area (ratio of i1 + i2 area:m1 area) (Ross et al. 1998 character i5)'
[5] 'i1:i2 proportions (Ross et al. 1998 character i6)'
[6] 'i2 cross-sectional shape (ratio of mesiodistal length to buccolingual breadth) (Ross et al. 1998 character i8)'
[7] 'lower incisor crown height (Ross et al. 1998 character i9)'
[8] 'lower incisor root orientation (Ross et al. 1998 character i11)'
[9] 'incisor crown orientation (Ross et al. 1998 character i12)'
[10] 'first lower incisor crown shape (Ross et al. 1998 character i14)'
[11] 'i2 heel development (Ross et al. 1998, character i15)'
[12] 'lower first incisor lingual cingulum (Ross et al. 1998 character i17)'
[13] 'i1 to m1 area (Ross et al. 1998 character i19)'
[14] 'female canine maximum cross-sectional area relative to first lower molar area (Ross et al. 1998 character c1, modified)'
[15] 'canine dimorphism (square root of male lower canine area:square root of female lower canine area) (Ross et al. 1998 character c2)'
[16] 'lower canine cross-sectional shape (Ross et al. 1998 character c3)'
[17] 'orientation of lower canine root'
[18] 'lower canine lingual crest development (Ross et al. 1998 character c4)'
[19] 'lower canine paracristid (Ross et al. 1998 character c5, modified)'
[20] 'lower canine height (Ross et al. 1998 character c6, modified)'
[21] 'p1 presence (Ross et al. 1998 character p1, modified)'
[22] 'p2 roots presence (Ross et al. 1998 character p3, modified)'
[23] 'p2 metaconid (Ross et al. 1998 character p15)'
[24] 'p2 protoconid height and shape (Ross et al. character p13)'
[25] 'p2 buccal cingulum development (Ross et al. character p28)'
[26] 'p3 paraconid (Ross et al. 1998 character p6, modified)'
[27] 'p3 lingual cingulum'
[28] 'p3 metaconid size (Ross et al. 1998 character p16)'
[29] 'p3 lateral protocristid orientation (Ross et al. 1998, character p22, modified)'
[30] 'p3 roots (Ross et al. 1998 character p4, in part, modified)'
[31] 'p3 root orientation (contains information similar to Ross et al. 1998 character p45)'
[32] 'p3 hypoconid (contains information similar to that of Ross et al. 1998 character p25)'
[33] 'p3 entoconid and lingual talonid crest (Ross et al. 1998 character p19)'
[34] 'p3-4 protoconid height (Ross et al. 1998 character p41)'
[35] 'p3 area:p4 area (Ross et al. 1998 character p42)'
[36] 'p4 root number (Ross et al. 1998 character p4, in part)'

[37] 'p4 metaconid size (Ross et al. 1998 character p17)'

[38] 'spacing of p4 metaconid and protoconid (Ross et al. 1998 character p14)'

[39] 'mesiodistal position of p4 metaconid with respect to protoconid'

[40] 'p4 paraconid size (Ross et al. 1998 character p7, modified)'

[41] 'p4 paraconid position (Ross et al. 1998 character p9)'

[42] 'p4 lateral protocristid morphology (contains information similar to Rp21-42 and Rp23-44)'

[43] 'p4 premetacristid (Ross et al. 1998 character p18, modified)'

[44] 'p4 postprotocristid (Ross et al. 1998 character p36)'

[45] 'p4 postmetacristid (contains information similar to Ross et al. 1998 character p37)'

[46] 'p4 entoconid and lingual talonid crest (Ross et al. 1998 character p20)'

[47] 'height of p4 talonid'

[48] 'p4 cristid obliqua development (Ross et al. 1998 character p11)'

[49] 'position of p4 hypoconid or distal terminus of p4 cristid obliqua (Ross et al. 1998 character p26, modified)'

[50] 'p4 hypocristid shearing development (Ross et al. 1998 character p27)'

[51] 'lower premolar crowding (Ross et al. 1998 character p5)'

[52] 'lower premolar inflation (Ross et al. 1998 character p29)'

[53] 'premolar orientation (Ross et al. 1998 character p33)'

[54] 'p4 exodaenodonty (Ross et al. 1998 character p30)'

[55] 'buccal cingulum on p4 (Ross et al. 1998 character p34)'

[56] 'p4 lingual cingulum'

[57] 'p4 md length:bl width (Ross et al. 1998 character p43)'

[58] 'p4 area relative to m1 area (Ross et al. 1998 character p44)'

[59] 'm1 length:m1 width (Ross et al. 1998 character m55)'

[60] 'm1 area (Ross et al. 1998 character m54)'

[61] 'm1 paraconid size (contains information similar to Ross et al. 1998 character m20)'

[62] 'm1 paraconid position (Ross et al. 1998 character m8)'

[63] 'm1 parastylid (Ross et al. 1998 character m11)'

[64] 'm1 premetacristid (contains information similar to Ross et al. 1998 character m18)'

[65] 'm1 metaconid position (Ross et al. 1998 character m19)'

[66] 'm1 trigonid height (ratio of trigonid height to talonid height) (Ross et al. 1998 character m16)'

[67] 'm1 cristid obliqua orientation (Ross et al. 1998 character m32)'

[68] 'm1 cristid obliqua terminus (Ross et al. 1998 character m34)'

[69] 'hypoconulid position on m1 (contains information similar to Ross et al. 1998 character m30)'

[70] 'm1 hypoconulid size (Ross et al. 1998 character m27)'

[71] 'm1 cristid obliqua development (Ross et al. 1998 character m31, in part)'

[72] 'm1-2 hypocristid development (Ross et al. 1998 character m38)'

[73] 'm1-2 buccal cingulum development (Ross et al. 1998 character m45)'

[74] 'm1-2 talonid -- lingual configuration (Ross et al. 1998 character m40)'

[75] 'm2 paraconid size (contains information similar to Ross et al. 1998 character m20)'

[76] 'm2 paraconid position (Ross et al. 1998 character m9)'

[77] 'm2 trigonid width (ratio of buccolingual breadths of trigonid and talonid) (Ross et al. 1998 character m6)'

[78] 'm2 cristid obliqua orientation (Ross et al. 1998 character m33)'

[79] 'm2 cristid obliqua terminus (Ross et al. 1998 character m35)'

[80] 'm2 cristid obliqua development (Ross et al. 1998 character m31, in part)'

[81] 'position of entoconid relative to hypoconid on m2 (contains information similar to Ross et al. 1998 character m57)'
[133] 'P3-4 buccal cingulum development (RP21-151)'
[134] 'upper premolar paraconules (contains information similar to RP14-144)'
[135] 'P4 lingual cingulum (contains information similar to RP19-149)'
[136] 'P4 metacone (RP10-140)'
[137] 'P4 protocone (RP11-141)'
[138] 'P4 hypocone (RP13-143, in part)'
[139] 'P4 pericone'
[140] 'P4 metaconules'
[141] 'P4 postprotocrista (RP17-147, in part)'
[142] 'P4 occlusal outline (RP7-137)'
[143] 'P4 root number (RP3-133)'
[144] 'P4 area:M1 area (RP5-135)'
[145] 'M1 pericone (RM20-170, modified)'
[146] 'M1 area:M2 area (RM4-155)'
[147] 'M1 mesiobuccal fovea delimited by trenchant parahypocrista'
[148] 'M1-2 paraconule size (RM37-180)'
[149] 'M1-2 paraconule position (RM15-165)'
[150] 'M1 hypocone size (RM10-160)'
[151] 'M1-2 hypocone position (RM12-162)'
[152] 'convex distal lobe for M1-2 hypocone'
[153] 'lingual extension of M1 hypocone lobe'
[154] 'M2 prehypocrista development (RM13-163)'
[155] 'M1-2 Nannopithex fold (RM5-156)'
[156] 'M1-2 postprotocrista development (RM18-168)'
[157] 'M1-2 metaconule size (RM16-166)'
[158] 'M1-2 premetaconule cristae (RM27-173)'
[159] 'M1-2 postmetaconule cristae (RM28-174)*'
[160] 'M1-2 lateral posterior transverse crista development (RM19-169)'
[161] 'shape of M2 centrocrista'
[162] 'buccal cingulum on M1-2 (contains information similar to RM24-172)'
[163] 'M1-2 root number (RM1-152)'
[164] 'M2 shape (b1:md) (RM3-154)'
[165] 'M2 pericone'
[166] 'M2 hypocone size (RM11-161)'
[167] 'M2 postmetacrista'
[168] 'M2 buccal expansion of paracone (RM33-177)'
[169] 'M1-3 lingual cingulum development (RM22-171)'
[170] 'M1-3 anterior cingulum (RM44-181)'
[171] 'molar protocone lingual inflation (RM31-176)'
[172] 'M3 metacone (RM34-178)'
[173] 'M3 paraconule (RM30-175)'
[174] 'M3 hypocone (RM36-179)'
[175] 'M3 size relative to M1 (RM46-182)'
[176] 'M3 root number (RM2-153)'
[177] 'mandibular symphyseal fusion (Y57,RCr39-221)'
[178] 'shape of the mandibular angle (Y40)'
[179] 'shape of masseteric fossa'
[180] 'mandibular depth (Y39,RCr45-227)'
[181] 'height of coronoid process relative to condyle (RCr43-225)'
[182] 'condyle height relative to toothrow (RCr44-226)'
[183] 'scapular morphology (mediolateral width / inferosuperior height) (contains information similar to that of Y64)'
[184] 'length of scapular coracoid process (coracoid process length / glenoid fossa height)'
[185] 'relative sizes of infraspinous and supraspinous fossae (Y66)'
[186] 'position of humeral head relative to tubercles'
[187] 'morphology of deltopectoral crest (RH16-247)'
[188] 'shape of intertubercular sulcus (contains information similar to RH15-246)'
[189] 'teres major insertion'
[234] 'bony distinction between plantar edge of ectal facet and body'

[235] 'size of calcaneal peroneal tubercle (100\*widest point on peroneal tubercle to ectal facet, measured perpendicular to long axis of bone / calcaneal width)'

[236] 'peroneal tubercle shape'

[237] 'number of "peaks" on calcaneal peroneal tubercle'

[238] 'position of peroneal process relative to sustentaculum'

[239] 'position of the peroneal tubercle relative to ectal facet (RC2-276)'

[240] 'development of distal plantar tubercle on calcaneus'

[241] 'level of distal plantar tubercle (tubercle to cuboid facet:calcaneal length)'

[242] 'orientation of long axis of calcaneocuboid joint'

[243] 'calcaneocuboid joint shape'

[244] 'concavity of cuboid facet'

[245] 'dorsal extension of bone supporting cuboid facet'

[246] 'posterior (plantar) calcaneal bowing'

[247] 'mediolateral bowing of calcaneal tuber'

[248] 'calcaneal sustentacular facet configuration'

[249] 'form of medial margin between sustentaculum and anterior calcaneal facet (viewed dorsally)'

[250] 'facet anterior to the anterior extension of the sustentacular facet (navicular or otherwise)'

[251] 'angle between sustentaculum and ectal facet'

[252] 'shape of medial edge of sustentaculum in plantar view'

[253] 'morphology of groove for flexor fibularis under sustentaculum'

[254] 'position of distal terminus of ectal facet relative to anterior calcaneal segment'

[255] 'size of tuber calcanei'

[256] 'shape of tuber calcanei'

[257] 'dorsal heel process'

[258] 'navicular length relative to width (contains information similar to RN1-278)'

[259] '"three-pronged" distal navicular articulation'

[260] 'morphology of the naviculocuboid articulation (RN3-279)'

[261] 'MTI relative dorsoplantar diaphysis midshaft breadth'

[262] 'MTI proximal articular surface angle, in dorsolateral view with respect to the proximal end of the bone'

[263] 'MTI proximal articular surface shape in dorsolateral view'

[264] 'MTI proximal articular surface orientation in proximal view (i.e., torsion)'

[265] 'MTI peroneal process proximodistal length in dorsal view with respect to the distal end of the bone'

[266] 'MTI peroneal process dorsoplantar thickness in dorsolateral view with respect to the proximal end of the bone'

[267] 'MTI medial side of proximal articular surface with respect to the proximal end of the bone'

[268] 'MTI dorsal notch located proximal to the distal articular surface in dorsal or lateral view'

[269] 'MTI head distal projection in dorsal view'

[270] 'MTI head shape in distal view'

[271] 'MTI distal epicondyle asymmetry in distal and dorsal views'

[272] 'MTI dorsoplantar axis of head orientation relative to long axis of diaphysis in plantar view, with respect to the distal end of the bone'

[273] 'MTI proximal articular surface notch location'

[274] 'MTI distal articular surface midline keel'

[275] 'MTI medial proximally projecting tubercle size'

[276] 'MTI dorsal extension of proximal articular surface'

[277] 'lateral process of entocuneiform (RE2-281)'

[278] 'foot axis (R01-282)'

[279] 'metatarsus length (MT3 length / proximal phalanx of digit III) (contains information similar to that of R04-285)'
[280] 'length of MT1 relative to MT3 (RMT2)'

[281] 'tibial process for peroneus longus'

[282] 'retroflexion of proximal tibial articulation'

[283] 'fusion of tibia and fibula (RT1-263)'

[284] 'shape of distal surface of tibia (RT3-264)'

[285] 'rotation of the medial malleolus (RT4-265)'

[286] 'shape of the medial malleolar articular surface (RT5-266)'

[287] 'shape of distal tibial shaft (RT6-267)'

[288] 'position of tibialis posterior groove (RT7-268)'

[289] 'fovea capitis (Y75)'

[290] 'femoral head shape (RF6-256)'

[291] 'morphology of femoral head articular surface'

[292] 'femoral neck length'

[293] 'angle of femoral neck (RF2-252)'

[294] 'anteroposterior angulation of proximal femur (contains information similar to RF8-258)'

[295] 'triangular depression on medial side of proximal femur (Dagosto and Schmid, 1996)'

[296] 'greater trochanter shape (Y72)'

[297] 'crista paratrochanterica'

[298] 'relative length of trochanteric fossa (RF9-259)'

[299] 'intertrochanteric crest (RF10-260)'

[300] 'angle of lesser trochanter (RF3-253)'

[301] 'gluteal tuberosity (Y74, contains information similar to RF4-254)'

[302] 'position of gluteal tuberosity'

[303] 'patellar margin asymmetry (raised and sharply defined lateral patellar margin) (contains information similar to RF12-262)'

[304] 'knee index (contains information similar to RF5-255)'

[305] 'shape of manubrium (Y83)'

[306] 'transpedicular foramina (Y85)'

[307] 'number of thoraco-lumbar vertebrae'

[308] 'anterior inferior iliac spine (Y70)'

[309] 'ilium shape (Y67)'

[310] 'ischial flaring (Y69)'

[311] 'ischium: ilium index (Anemone & Covert, 2000)'

[312] 'pubic bone flaring (Y68)'

[313] 'tail length (Y86)'

[314] 'shape of glenoid fossa (RCr40-222)'

[315] 'postglenoid process (Y25)'

[316] 'postglenoid process-auditory bulla relationship (Y26)'

[317] 'postglenoid foramen'

[318] 'position of postglenoid foramen relative to postglenoid process'

[319] 'entoglenoid process (RCr41-223)'

[320] 'position of pyramidal processes (RCr35-217)'

[321] 'position of the anteriormost point on the palatine / maxillary suture of the palate (Y19)'

[322] 'posterior palatine torus (RCr34-216)'

[323] 'maxillary-palatine notch'

[324] 'position of the posteromedial edge of the palate in relation to M3 (Y24)'

[325] 'shape of medial pterygoid plate (RCr36-218, contains information similar to Y22)'

[326] 'lateral pterygoid-bullar overlap (RCr18-200)'

[327] 'extent of contact between the lateral pterygoid plate and the bullar wall (RCr19-201)'

[328] 'position of foramen ovale (Y28)'

[329] 'central stem of basicranium (D47)'

[330] 'choanal shape (D50)'

[331] 'basioccipital flange (contains information similar to RCr20-202)'

[332] 'encroachment of auditory bulla on pterygoid fossa (RCr17-199)'

[333] 'shape of external auditory meatus (D38)'
[334] 'suprameatal foramen (RCr21-203)'

[335] 'ascending pharyngeal artery (Y52)'

[336] 'vascular plexus associated with ascending pharyngeal artery (Y53)'

[337] 'position of chorda tympani relative to m. tensor tympani (Maier, 2008)'

[338] 'relative sizes of stapedial canal (SC) and promontory canal (PC)'

[339] 'mediolateral position of posterior carotid foramen (RCr6-188)'

[340] 'rostrocaudal position of posterior carotid foramen with respect to fenestra cochleae (contains information similar to RCr8-190)'

[341] 'ventrodorsal position of posterior carotid foramen relative to fenestra cochleae (RCr7-189)'

[342] 'position of pathway for internal carotid artery or nerve relative to fenestra cochleae (RCr9-191)'

[343] 'morphology of path for promontory artery and / or internal carotid nerve (RCr12-194)'

[344] 'perbullar pathway for internal carotid artery (RCr4-186)'

[345] 'transverse septum defining caudal wall of anterior accessory cavity (contains information similar to RCr1-183)'

[346] 'trabeculated anterior accessory cavity'

[347] 'number of cochlear turns'

[348] 'composition of bulla'

[349] 'epitympanic crest (RCr48-230)'

[350] 'morphology of annular bridge (RCr16-198)'

[351] 'position of ventral edge of tympanic bone (RCr14-196)'

[352] 'parotic fissure (RCr22-204)'

[353] 'pneumatization of mastoid (from epitympanic recess) (RCr3-185)'

[354] 'paroccipital processes (Y23)'

[355] 'emissary foramina on lateral edge of parietals (Y33)'

[356] 'pattern of sagittal cresting'

[357] 'metopic sutural fusion in adults (RCr31-213)'

[358] 'frontal-maxillary contact'

[359] 'degree of orbital convergence (contains information similar to RCr32-214)'

[360] 'expansion of ethmoturbinals (YIP89)'

[361] 'transverse lamina:ethmoturbinal recess'

[362] 'pronounced interorbital constriction (RCr27-209)'

[363] 'orbit size (contains information similar to RCr23-205)'

[364] 'foramen rotundum (Y12,RCr29-211)'

[365] 'lacrimal foramen morphology (Y11)'

[366] 'position of lacrimalforamen (RCr30-212)'

[367] 'zygomatic-lacrimal contact (Y6, RCr26-208)'

[368] 'lacrimal-palatine contact (RCr28-210)'

[369] 'ethmomaxillary fissure (Y13, modified following Cartmill 1978)'

[370] 'anterior and posterior hiatus of ethmomaxillary fissure (Cartmill, 1978)'

[371] 'frontal-palatine contact (Cartmill, 1978)'

[372] 'position of sphenopalatine foramen'

[373] 'position of infraorbital foramina (contains information similar to Y3)'

[374] 'morphology of zygomatic portion of postorbital bar (Y18, contains information similar to RCr24-25)'

[375] 'morphology of frontal segment of the postorbital bar (Y17, contains information similar to RCr24-25)'

[376] 'zygomatic arch depth'

[377] 'zygomaticofacial foramen (Y16)'

[378] 'snout length (RCr37-219)'

[379] 'maxillary depth (RCr38-220)'

[380] 'facial profile (Masters & Brothers, 2002)*'

[381] 'position of anterior palatine foramina relative to incisors (Masters & Brothers, 2002)*'

[382] 'rostral projection of premaxilla (Masters & Brothers, 2002)*'
[383] 'ascending wing of premaxilla (RCr49-231)'
[384] 'retia mirabilia of the proximal limb vessels (YIP87)'
[385] 'nasal morphology (Y62, contains information similar to R289)'
[386] 'retinal fovea (Y60, R288)'
[387] 'tapetum lucidum (Y59, R290)'
[388] 'presence-absence of choriovitelline placenta (Y58, KPL4)'
[389] 'epitheliochorial vs. hemochorial placentation (KPL1)'
[390] 'blastocyst attachment (KPL2)'
[391] 'amniotic cavity (KPL3)'
[392] 'embryonic body stalk (KPL5)'
[393] 'allantois development (KPL6)'
[394] 'ascorbic acid synthesis'
;
STATELABELS

1
'three incisors in each quadrant '
'two incisors in each quadrant '
'one incisor in each quadrant '
'no incisors present ',

2
'arcuate battery in occlusal view ("U"-shaped arcade) '
'incisor cusp tips staggered ("V"-shaped arcade) ',

3
'lower canine is not incorporated into a toothcomb '
'lower canine is incorporated into a toothcomb ',

4
'small (<=0.69) '
'intermediate (>=0.70, <=1.07) '
'large (>1.07) ',

5
'i1 / i2 area ratio <0.65 '
'0 / 2 '
'i1 / i2 area ratio =>0.65, <0.82 '
'2 / 4 '
'i1 / i2 area ratio =>0.83, <1.00 '
'4 / 6 '
'i1 / i2 area ratio =>1.01, <1.25 '
'6 / 8 '
'i1 / i2 area ratio =>1.25 ',

6
'ratio =>0.64 '
'ratio <=0.64 ',

7
'low-crowned '
'0 / 2 '
'moderately highcrowned '
'2 / 4 '
'high-crowned ',

8
'erect or 'vertical '
'0 / 2 '
'slightly procumbent '
'2 / 4 '
'very procumbent ',

9
'erect or vertical '
'0 / 2 '
'procumbent '
'2 / 4 '
'very procumbent ',

10
'spatulate '
'pointed or lanceolate ',

11

'heel absent '
'heel present ',

12

'absent to weak '
'0 / 2 '
'strong but incomplete '
'2 / 4 '
'strong and complete ',

13

'i1 area << m1 area '
'0 / 2 '
'i1 area < or = m1 area '
'2 / 4 '
'i1 > m1 ',

14

'c1 area / m1 area < 0.40 '
'0 / 2 '
'c1 area / m1 area =>0.4, <0.8 '
'2 / 4 '
'c1 area / m1 area =>0.8, <=1.2 ',

15

'<1.07 '
'>=1.07, <1.17 '
'>=1.17 ',

16

'rounded oval '
'mesiodistally compressed '
'buccolingually compressed ',

17

'oriented in line with long axis of crown '
'mesially inflected ',

18

'rounded '
'0 / 2 '
'sharp ',

19

'oblique to occlusal plane '
'nearly horizontal to occlusal plane '
'forms part of cropping mechanism with i1-2 '
'oriented in line with buccal face of adjacent incisor ',

20

'low, squat '
'0 / 2 '
'narrow, short '
'tall, at or above toothrow '
'high crowned but very procumbent ',

21

'present '
'0 / 2 '
'absent ',

22

'two '
'0 / 2 '
'one '
'2 / 4'
'p2 absent ',

23
'metaconid absent or indistinct '
'0 / 2 '
'metaconid is small '
'2 / 4 '
'metaconid is large ',
24
'extremely short, shorter than p3 '
'0 / 2 '
'not projecting, in line with p3 '
'2 / 4 '
'slender, projects above protoconids of p3-4 '
'4 / 6 '
'massive, projects above protoconids of p3-4 ',
25
'absent '
'0 / 2 '
'incomplete, broken at protoconid and hypoconid '
'2 / 4 '
'complete ',
26
'present, large '
'0 / 2 '
'present, small '
'2 / 4 '
'negligible or absent ',
27
'present '
'0 / 2 '
'absent ',
28
'absent '
'0 / 2 '
'present, small '
'2 / 4 '
'present, large ',
29
'transversely oriented '
'0 / 2 '
'distally or distolingually oriented '
'2 / 4 '
'absent ',
30
'one '
'0 / 2 '
'two ',
31
'mesial root placed directly mesial to distal root'
'0 / 2 '
'mesial root placed lateral to distal root',
32
'large and well-defined, projects above other talonid cusps'
'0 / 2 '
'very small, absent, or incorporated into talonid as an indistinct cusp',
33
absent
'0 / 2 '
'lingual talonid crest present but an entoconid does not stand out above it'
'2 / 4 '
'entoconid is a small discrete cusp',

34

'p3 much lower than p4'

'0 / 2'

'p3 slightly lower than p4'

'2 / 4'

'p3 equal in height to p4'

'4 / 6'

'p3 higher than p4',

35

'<0.45'

'0 / 2'

'0.45-0.59'

'2 / 4'

'0.60-0.69'

'4 / 6'

'0.70-0.79'

'6 / 8'

>0.80',

36

one

two,

37

'absent or indistinct'

'0 / 2'

small

'2 / 4'

large,

38

'close to protoconid'

'0 / 2'

'widely spaced from protoconid',

39

'metaconid is distal to protoconid'

'0 / 2'

'metaconid is placed transverse to protoconid',

40

'p4 paraconid is present and large'

'0 / 2'

'p4 paraconid is present and small'

'2 / 4'

'p4 paraconid is very small or absent',

41

'mesial to protoconid'

'mesiolingual, between protoconid and metaconid'

'mesial to metaconid, widely spaced from metaconid',

42

'protocristid poorly developed or absent'

'0 / 2'

'distolingually oriented protocristid present'

'2 / 4'

'trenchant and tranversely oriented protocristid present',

43

'premetacristid absent or poorly developed'

'0 / 2'

'premetacristid present',

44

'weak or absent'

'0 / 2'

moderate

'2 / 4'
'very strong',

45

'weak or absent'
'0 / 2'
moderate
'2 / 4'
'very strong',

46

'absent or a trace'
'0 / 2'
'lingual talonid crest present but an entoconid does not stand out above it'
'2 / 4'
'entoconid is a small discrete cusp',

47

'no aspect of the talonid is at level of m1 trigonid basin'
'0 / 2'
'hypoconid extends to level of m1 trigonid basin'
'2 / 4'
'entire talonid basin extends to level of m1 trigonid basin',

48

absent
'0 / 2'
weak
'2 / 4'
strong,

49

'buccal to protoconid'
'0 / 2'
'distal to protoconid'
'2 / 4'
'between protoconid and metaconid'
'4 / 6'
'distal to metaconid',

50

absent
'0 / 2'
weak
'2 / 4'
strong,

51

'no crowding'
'0 / 2'
'slightly crowded'
'2 / 4'
'very crowded',

52

'not basally inflated'
'0 / 2'
'slightly basally inflated'
'2 / 4'
'very basally inflated',

53

'crown bases vertical in lateral perspective'
'0 / 2'
'slightly oblique'
'2 / 4'
'strongly oblique',
'projecting mesially',

54

'not exodaenodont'
'slightly exodaenodont'
'very exodaenodont',

55

'absent or indistinct'
'0 / 2'
'present and distinct',

56

'absent or discontinuous'
'0 / 2'
present,

57

'<0.95'
'0 / 2'
'>0.96, <1.14'
'2 / 4'
'>1.15, <1.35'
'4 / 6'
'>1.36, <1.46'
'6 / 8'
'>1.47',

58

'<0.62'
'0 / 2'
'>0.63, <0.72'
'2 / 4'
'>0.73, <0.82'
'4 / 6'
'>0.83, <0.92'
'6 / 8'
'>0.93',

59

'1.0-1.15'
'0 / 2'
'1.16-1.22'
'2 / 4'
'1.23-1.32'
'4 / 6'
'>1.33',

60

'1.10-3.00 mm'
'0 / 2'
'3.10-5.00 mm'
'2 / 4'
'5.10-7.00 mm'
'4 / 6'
'7.10-9.00 mm'
'6 / 8'
'>9.10 mm',

61

'absent or crestiform'
'0 / 2'
'present, small'
'2 / 4'
'present, large',

62

'mesial to protoconid'
'mesiolingual, equidistant between protoconid and metaconid'
'1 / 3'
'mesial to metaconid but widely spaced from it twinned with
metaconid',

63
absent
'0 / 2'
present,

64

'premetacristid absent or indistinct'
'0 / 2'
'premetacristid present,

65

'approximately transverse to protoconid'
'0 / 2'
'distal to protoconid',

66

'higher than talonid (=>1.20)'
'0 / 2'
'slightly higher than talonid (=>1.10, <1.20)'
'2 / 4'
'trigonid and talonid of similar height',

67

'reaches trigonid wall at a point distal to protoconid'
'0 / 2'
'reaches trigonid wall at a point distolingual to protoconid'
'2 / 4'
'reaches trigonid wall at a point distal to metaconid',

68

'runs to base of trigonid'
'0 / 2'
'runs part way up the distal trigonid wall'
'2 / 4'
'connects with protoconid tip or protocristid'
'4 / 6'
'connects with metaconid',

69

'twinned with, or most closely situated next to, entoconid'
'0 / 2'
central
'2 / 4'
'closer to hypoconid than to entoconid',

70

large
'0 / 2'
moderate
'2 / 4'
small
'4 / 6'
absent,

71

'weak (rounded)'
'0 / 2'
'strong (trenchant)'

72

'absent or indistinct'
'0 / 2'
weak
'2 / 4'
strong,

73

'absent to trace'
'0 / 2'
'partial, discontinuous'
'2 / 4'
complete,
74
open
'0 / 2'
'notched lingually or closed',

75
absent
'0 / 2'
'present, small'
'2 / 4'
'present, large',

76
'mesial to protoconid'
'0 / 2'
'mesiolingual,
'approximately equidistant between protoconid and metaconid'
'2 / 4'
'mesial to metaconid but widely spaced from it twinned with
metaconid',

77
'much wider than talonid (=>1.11)'
'0 / 2'
'width similar (<1.11,>0.90)'
'2 / 4'
'much narrower than talonid (=<0.90)',

78
'reaches trigonid wall at a point distal to protoconid '0 / 2'
'reaches trigonid wall at a point distolingual to protoconid'
'2 / 4'
'reaches trigonid wall at a point distal to metaconid',

79
'runs to base of trigonid'
'0 / 2'
'runs part way up distal trigonid wall'
'connects to protoconid tip or protocristid
'3 / 5'
'connects with metaconid',

80
'weak (rounded)'
'0 / 2'
'strong (trenchant)',

81
'mesial to hypoconid'
'0 / 2'
'transverse to hypoconid'
'2 / 4'
'distal to hypoconid',

82
'twinned with, or most closely situated next to, entoconid'
'0 / 2'
'central
'2 / 4'
'closer to hypoconid than to entoconid',

83
large
'0 / 2'
'moderate
'2 / 4'
'small
'4 / 6'
'absent,
84
'no postentoconid sulcus'
'hypoconulid closely situated next to entoconid and postentoconid sulcus present, entoconid confluent with distal talonid wall'
'distolingual fovea present, entoconid confluent with posterior talonid wall'

85
'distolingual fovea present, entoconid mesiodistally abbreviated and not confluent with posterior talonid wall',

absent
'variously present',

86
crestiform
'0 / 2'
'moderate basal inflation'
'2 / 4'
'cusp bases bulbous',

87
'm3 much longer than m2 (0.71-0.80)'
'0 / 2'
'm3 longer than m2 (0.81-0.90)'
'2 / 4'
'm3 slightly longer or equal in length to m2 (0.91-1.00)'
'4 / 6'
'm3 slightly shorter than m2 (1.01-1.12)'
'6 / 8'
'm3 much shorter than m2 (>=1.13) or absent',

88
absent
'0 / 2'
'present, small'
'2 / 4'
'present, large',

89
'mesial to protoconid'
'0 / 2'
'mesiolingual, between protoconid and metaconid'
'2 / 4'
'mesial to metaconid but widely spaced from it'
'4 / 6'
'twinned with metaconid',

90
'absent or indistinct'
'runs to base of trigonid runs part way up distal trigonid wall'
'connects with protoconid tip or protocristid connects with metaconid',

91
'=>1.20'
'0 / 2'
'<1.20->1.05'
'2 / 4'
'<1.05',

92
'present and distinct'
'0 / 2'
'greatly reduced or absent',

93
lobate
'0 / 2'
'not lobate',

94
'centrally located'

'twinned with, or situated closest to, entoconid'

'buccally located, closer to hypoconid',

95

absent

'0 / 2'

single

'2 / 4'

'double, or additional cusp distal to entoconid that is independent

of distal lobe',

96

one

two,

97

absent

present,

98

absent

'0 / 2'

'present, protocristids poorly developed or absent',

99

absent

'0 / 2'

'trenchant postmetacristid or incipient metastylid'

'2 / 4'

'present, large',

100

smooth

'0 / 2'

'slightly crenulated'

'2 / 4'

'highly crenulated',

101

'very broad'

narrow,

102

'absent, teeth widely spaced'

'0 / 2'

'present as a narrow contact'

'2 / 4'

'I2 tightly packed against I1, I1 paracrista abbreviated',

103

'areas approximately equal or I1 smaller than I2 (<=1.00)'

'0 / 2'

'I1 slightly larger than I2 (>1.00,<1.40)'

'2 / 4'

'I1 much larger than I2 (>1.40)',

104

'weak, discontinuous'

'0 / 2'

'moderate, continuous'

'2 / 4'

strong,

105

absent

present,

106

'rounded oval (<1.05)'

'0 / 2'

'buccolingually compressed (>1.05, <1.30)'

'2 / 4'
'extremely compressed (>1.30)',
107
'spatulate (no apparent occlusal cusp, mesial and distal edges
continuous and rounded)'
'0 / 2'
'semi-spatulate (central cusp present, but blunt with discernable
mesial and distal occlusal crests)'
'2 / 4'
'central occlusal cusp pointed, occlusal edges steep',
108
'small (<=0.50)'
'moderate (>0.5, <0.56)'
'large (>0.56)',
109
'occlusal edge approximately orthogonal to long axis of root'
'occlusal edge wears at a steep angle to long axis of root'
'crown with pronounced mesial asymmetry (=mesial process) in unworn
state',
110
absent
present,
111
'rounded oval (<=1.05)'
'0 / 2'
'slightly buccolingually compressed (>1.05, <1.30)'
'2 / 4'
'extremely buccolingually compressed (>=1.30)',
112
present
'0 / 2'
absent,
113
'three in each quadrant'
'two in each quadrant'
'one or none in each quadrant',
114
'round in cross-section'
'0 / 2'
oval
'2 / 4'
'strongly compressed buccolingually',
115
'long axis of canine oriented in line with adjacent premolar'
'canine is slightly rotated internally'
'canine is strongly rotated internally',
116
'shallow or absent'
deep,
117
'weak or absent'
'0 / 2'
strong
'2 / 4'
'very strong',
118
'wears against p1-2'
'0 / 2'
'wears against p2'
'2 / 4'
'wears against p2-3'
'4 / 6'
'wears against p3',
119
  present
  absent,
120
  three
  two
  one
  P2 absent
121
  triangular
  'suboval with the long axis buccolingual'
  'suboval with the long axis mesiodistal'
  round,
122
  present
  absent,
123
  'smaller than, or subequal in height (P2 height / P3 height <=1.1)'
  '0 / 2'
  'P2 slightly taller than P3 (P2 height / P3 height >=1.11, <1.5)'
  '2 / 4'
  'P2 much taller than P3 (P2 height / P3 height >1.51)',
124
  'much smaller than P3 (<=0.85) (if P2 is absent, taxon is assigned state 0)'
  '0 / 2'
  'approximately the same size, or only slightly smaller, than P3 (>0.85, <0.95)'
  '2 / 4'
  'much larger than P3 (>=.95)',
125
  'smoothly rounded'
  '0 / 2'
  'waisted between buccal and lingual cusps',
126
  absent
  '0 / 2'
  'present, small'
  '2 / 4'
  'present, distinct',
127
  absent
  '0 / 2'
  present,
128
  absent
  '0 / 2'
  present,
129
  one
  '0 / 2'
  two
  '2 / 4'
  three,
130
  present
  '0 / 2'
  absent,
131
  absent
'0 / 2'
present,

132
'trigon>=talon'
'trigon<talon',

133
'absent or weak'
'0 / 2'
strong,

134
'absent or indistinct'
'transverse crest connecting protocone to paracone'
present,

135
absent
'0 / 2'
present,

136
absent
'0 / 2'
present,

137
'low relative to paracone'
'high relative to paracone',

138
absent
'0 / 2'
present,

139
absent
present,

140
absent
'0 / 2'
present,

141
absent
'0 / 2'
'weak, short'
'2 / 4'
strong
'4 / 6'
'strong, secondary postprotocrista directed toward metacone',

142
triangular
suboval
'1 / 3'
squared,

143
one
two
three,

144
'P4<<M1 (<=0.66)'
'0 / 2'
'P4<M1 (>0.66,<=0.76)'
'2 / 4'
'P4=M1 (0.77-1.05)'
'4 / 6'
'P4>M1 (>1.06)'

145
absent
'0 / 2'
small
'2 / 4'
large,

146
'M1>M2 (<1.40,>1.0)'
'0 / 2'
'M1=<M2 (<=1.0)',

147
absent
'0 / 2'
present,

148
absent
'0 / 2'
'small-moderate (smaller than paracone)'
'2 / 4'
'large (nearly as large as protocone)',

149
'attached to preprotocrista'
'0 / 2'
'not attached to preprotocrista',

150
large
'0 / 2'
small
'2 / 4'
absent,

151
'distal, slightly lingual to protocone'
'0 / 2'
'distal, far lingual to protocone',

152
absent
'0 / 2'
'present, distal crown margin weakly concave'
'2 / 4'
'present, distal crown margin deeply notched',

153
'lobe does not extend lingually far beyond
protocone'

154
'0 / 2'
'lobe does extend far lingually past protocone',
absent
'0 / 2'
weak
'2 / 4'
'strong -- reaches to postprotocrista, encloses the
talon lingually',

155
absent
'0 / 2'
weak
'2 / 4'
'strong ',

156
'strong, runs to base of metaconule or metacone'
'0 / 2'
'strong but short, does not reach to base of
metacone'
absent,
absent
'0 / 2'
small
'2 / 4'
moderate
'4 / 6'
large,
'absent or weak '
'0 / 2'
'strong ',
'absent or weak'
'0 / 2'
strong,
sharp
'0 / 2'
indistinct,
'straight, aligned with mesiodistal plane'
'0 / 2'
'crests are more buccally oriented, meet at an
'2 / 4'
'crests meet at a sharp angle or form a mesostyle',
absent
'0 / 2'
'present but poorly developed'
'2 / 4'
'present and distinct'
'4 / 6'
'extensive buccal "shelf",
'three or more, three or more'
'three, two',
'very broad (>1.65)'
'0 / 2'
'broad (<1.65,>1.30)'
'2 / 4'
'squared (<=1.30)',
absent
'0 / 2'
present,
large
'0 / 2'
small
'2 / 4'
absent,
'short, indistinct, or absent'
'long, trenchant, and labially extended',
'no expansion'
'0 / 2'
'slight expansion'
'2 / 4'
'considerable expansion',
169
absent
'0 / 2'
'weak, broken'
'2 / 4'
'strong, complete ',
170
'strong, complete, long'
'0 / 2'
'strong, short'
'2 / 4'
'weak or absent',
171
'not inflated'
'0 / 2'
'slightly inflated'
'2 / 4'
'very inflated',
172
'absent or very small'
'0 / 2'
'moderate (but smaller than paracone)'
'2 / 4'
'large (equal to paracone)',
173
absent
'0 / 2'
'small-moderate'
'2 / 4'
large,
174
'absent or very small'
'0 / 2'
small
'2 / 4'
large,
175
'very small (half the size of M1 or less)'
'0 / 2'
'small (two thirds)'
'2 / 4'
'large (approximately as large)',
176
three
'0 / 2'
two
'2 / 4'
one,
177
absent
'0 / 2'
present,
178
'sharply hooked '
'0 / 2'
'smoothly rounded '
'2 / 4'
'rounded but expanded posteriorly',
179 shallow
'0 / 2'
'deeply excavated',
180 'shallow (less than 1.8 times as deep @ m2 as md
length of m2)'
'0 / 2'
'deep (>1.8 times as deep @ m2 as md length of m2,
<2.19)'
'2 / 4'
'very deep (more than 2.2 times as deep @ m2 as md
length of m2)',
181 'very high above'
'slightly above or equal',
182 'at level of toothrow'
'slightly above'
'well above toothrow',
183 '<1.25'
'0 / 2'
'<1.5, >1.25'
'2 / 4'
'>1.5, <1.75'
'4 / 6'
'>1.75',
184 '<=.90 '
'0 / 2'
'>.9, <=1.0'
'2 / 4'
'>1.0, <=1.1 '
'4 / 6'
'>1.1',
185 'supraspinous fossa larger than infraspinous fossa '
'0 / 2'
'fossae roughly equivalent '
'2 / 4'
'infraspinous fossa approximately two times as large
as thesupraspinous fossa '
'4 / 6'
'infraspinous fossa approximately three times as
large as thesupraspinous fossa ',
186 'humeral head is expanded dorsal to tubercles'
'0 / 2'
'humeral head and tubercles are approximately equal
in height'
'2 / 4'
'tubercles are expanded above humeral head',
187 prominent
'0 / 2'
low,
188 'narrow and deep'
'0 / 2'
<table>
  <tr>
    <td></td>
    <td>'narrow and shallow'<br>'2 / 4'<br>'shallow and wide',</td>
  </tr>
  <tr>
    <td>189</td>
    <td>'absent or poorly defined'<br>'0 / 2'<br>'elongate well-defined crest'<br>'2 / 4'<br>'well-developed tubercle',</td>
  </tr>
  <tr>
    <td>190</td>
    <td>prominent<br>low,</td>
  </tr>
  <tr>
    <td>191</td>
    <td>broad<br>'0 / 2'<br>moderate<br>'2 / 4'<br>narrow,</td>
  </tr>
  <tr>
    <td>192</td>
    <td>'foramen is placed far proximal to trochlea'<br>'lateral wall of medial strut defining foramen is confluent with medial edge of the trochlea'<br>'0 / 2'<br>'foramen more medial, and lateral wall of medial strut defining foramen is not confluent with medial edge of the trochlea'<br>absent,</td>
  </tr>
  <tr>
    <td>193</td>
    <td>parallel<br>'0 / 2'<br>'slight dorsal angle'<br>'2 / 4'<br>'large dorsal angle',</td>
  </tr>
  <tr>
    <td>194</td>
    <td>reduced<br>'0 / 2'<br>prominent,</td>
  </tr>
  <tr>
    <td>195</td>
    <td>present<br>'0 / 2'<br>'small, shallow'<br>'2 / 4'<br>'absent',</td>
  </tr>
  <tr>
    <td>196</td>
    <td>'cylindrical, distomedial aspect of trochlea is straight'<br>'0 / 2'<br>'conical, medial aspect of trochlea flares distally',</td>
  </tr>
  <tr>
    <td>197</td>
    <td>subequal<br>'0 / 2'<br>'medial edge more flared than lateral edge',</td>
  </tr>
  <tr>
    <td>198</td>
    <td>'no pronounced lips on dorsal trochlear edges'<br>'0 / 2'<br>'both medial and lateral edges pronounced'<br>'very pronounced lateral lip',</td>
  </tr>
  <tr>
    <td>199</td>
    <td>'trochlea and capitulum are confluent, with no distinct trochleocapitularridge'<br>'0 / 2'</td>
  </tr>
</table>
<table>
  <tr>
    <td>by a weakridge'</td>
    <td>'lateral aspect of trochlea is offset from capitulum<br>'2 / 4'<br>'lateral aspect of trochlea is separated from<br>capitulum by a deep gutter',</td>
  </tr>
  <tr>
    <td></td>
    <td>200<br>globular<br>'0 / 2'<br>ovoid,</td>
  </tr>
  <tr>
    <td></td>
    <td>201<br>'elongate and distinct'<br>'short or absent'<br>'1 / 3'<br>'proximodistally tall capitular flange is present',</td>
  </tr>
  <tr>
    <td></td>
    <td>202<br>'ventral articular width / capitular width >2.5'<br>'0 / 2'<br>'&lt;2.5, &gt;2.0'<br>'2 / 4'<br>'&lt;2.0',</td>
  </tr>
  <tr>
    <td>transparent'</td>
    <td>203<br>'deep, open and unossified'<br>'0 / 2'<br>'moderately deep, thinly ossified and nearly<br>'2 / 4'<br>'shallow, robustly ossified',</td>
  </tr>
  <tr>
    <td></td>
    <td>204<br>'&lt;59'<br>'0 / 2'<br>'60-69'<br>'2 / 4'<br>'70-79'<br>'4 / 6'<br>'80-89'<br>'6 / 8'<br>'90-100',</td>
  </tr>
  <tr>
    <td>hamate'</td>
    <td>205<br>'facet faces distally, no articulation with hamate'<br>'facet faces distoradially, articulation with<br>'centrale not present as an independent element',</td>
  </tr>
  <tr>
    <td></td>
    <td>206<br>'no ulnar-pisiform articulation'<br>'facet on pisiform for ulnar styloid process is<br>smaller than, or roughly equal in size to, that for triquetrum'<br>'facet on pisiform for ulnar styloid process is much<br>enlarged and deeply excavated',</td>
  </tr>
  <tr>
    <td></td>
    <td>207<br>'no discernable styloid process'<br>'present but only moderately developed'<br>'long and well-developed',</td>
  </tr>
  <tr>
    <td></td>
    <td>208<br>'of normal length'<br>'drastically reduced',</td>
  </tr>
  <tr>
    <td></td>
    <td>209<br>'0-15 degrees'<br>'0 / 2'<br>'16-30 degrees'<br>'2 / 4'<br>'&gt;31 degrees',</td>
  </tr>
</table>
210

'nail present'
'claw present ',

211

present
absent,

212

'<50'
'0 / 2'
'>50, <60'
'2 / 4'
'>60, <70'
'4 / 6'
'>70',

213

'<100'
'0 / 2'
'100-120'
'2 / 4'
'>120',

214

absent
'0 / 2'
present,

215

shallow
'0 / 2'
moderate
'2 / 4'
'deep, medially projecting',

216

'< -0.151'
'0 / 2'
'between -0.15 and 0'
'2 / 4'
'between 0 and 0.15'
'4 / 6'
'> 0.16',

217

> 1.55'
'0 / 2'
'between 1.56 and 1.60'
'2 / 4'
'>1.61',

218

'<-0.151'
'0 / 2'
'between -0.15 and 0'
'2 / 4'
'between 0 and 0.15'
'4 / 6'
'between 0.151 and 0.30'
'6 / 8'
'>0.301',

219

'<100'
'0 / 2'
'>100, <120'
'2 / 4'
'>120',

220
'<20 degrees'
'0 / 2'
'20-30 degrees'
'2 / 4'
'>'30 degrees',

221
'head width / head height x 100 <115'
'0 / 2'
'head width / head height x 100 >115, <130'
'2 / 4'
'head width / head height x 100 >130',

222
absent
present,

223
absent
'0 / 2'
'present, small'
'2 / 4'
'present, large',

224
'79 degrees or less'
'0 / 2'
'80-89 degrees'
'2 / 4'
'90-99 degrees'
'4 / 6'
'100-109 degrees'
'6 / 8'
'110 degrees or higher,

225
'poorly developed'
'0 / 2'
small
'2 / 4'
large,

226
'>' -0.2'
'0 / 2'
'between -0.21 and -0.4'
'2 / 4' 'between -0.41 and -0.6'
'4 / 6'
'>' -0.61',

227
'< 1.0'
'0 / 2'
'between 1.01 and 1.25'
'2 / 4'
'between 1.26 and 1.5'
'4 / 6'
'>' 1.5',

228
'<35'
'0 / 2'
'>'35, <40'
'2 / 4'
'>'40, <45'
'4 / 6'
'>'45, <50'
'6 / 8'
'>'50',
229

>30,<40'
'0 / 2'
'>41,<50'
'2 / 4'
'>51,<60'
'4 / 6'
'>61,<70'
'6 / 8'
'>71',

230

.10-.15'
'0 / 2'
'.16-.20'
'2 / 4'
'.21-.25'
'4 / 6'
'.26-.30'
'6 / 8'
'>.31',

231

<50'
'0 / 2'
'>50, <60'
'2 / 4'
'>60',

232

<120 degrees'
'>=120 degrees',

233

absent
intermediate
present,

234

'no well-defined border'
'well-defined border present',

235

'massive, extends far laterally (>=21)'
'medium (<21)'
'highly reduced or too small to measure',

236

'discrete tubercle with clearly defined borders'
'tubercle gradually slopes into body (substantially

longer proximodistally relative to mediolateral width)',

237

'single tubercle present
'two "peaks" present',

238

'below lower half'
'upper half',

239

'maximum width of peroneal tubercle is placed distal

to distal terminus of ectal facet

'maximum width of peroneal tubercle is placed approximately at the distal terminus of the ectal facet'

'maximum width of peroneal tubercle is placed proximal to the distal terminus of the ectal facet, closer to the distal terminus of the ectal facet than the proximal'

'maximum width of peroneal tubercle is placed proximal to distal terminus of the ectal facet, closer to the proximal terminus of the ectal facet than distal',
<table>
  <tr>
    <td>240</td>
    <td>'small, poorly developed'<br>'0 / 2'<br>'well-developed',</td>
  </tr>
  <tr>
    <td>241</td>
    <td>'distal, near cuboid facet (0-0.19)'<br>'more proximal (&gt;0.20)',</td>
  </tr>
  <tr>
    <td>242</td>
    <td>dorsoventral<br>oblique<br>mediolateral,</td>
  </tr>
  <tr>
    <td>243</td>
    <td>oval<br>'fan-shaped, notch is articular'<br>'fan-shaped, notch is non-articular'<br>'oval but longest in dorsoplantar direction',</td>
  </tr>
  <tr>
    <td>244</td>
    <td>'flat (&gt;=160'<br>'shallow (&gt;=150, &lt;160'<br>'deep (&lt;150)',</td>
  </tr>
  <tr>
    <td>245</td>
    <td>absent<br>present,</td>
  </tr>
  <tr>
    <td>246</td>
    <td>absent<br>'present, moderately developed'<br>'present, extreme bowing',</td>
  </tr>
  <tr>
    <td>247</td>
    <td>absent<br>'present, moderate bowing'<br>'present, extreme bowing',</td>
  </tr>
  <tr>
    <td>248</td>
    <td>'single continuous sustentacular facet present'<br>'0 / 2'<br>'separate anterior and posterior sustentacular<br>facets present',</td>
  </tr>
  <tr>
    <td>249</td>
    <td>'waisted into body'<br>'relatively linear medial margin',</td>
  </tr>
  <tr>
    <td>250</td>
    <td>prominent<br>'very small or absent',</td>
  </tr>
  <tr>
    <td>251</td>
    <td>'&lt;140'<br>'140-160'<br>'&gt;160-&lt;180'<br>'&gt;180',</td>
  </tr>
  <tr>
    <td>252</td>
    <td>'triangular or beak-like'<br>rounded,</td>
  </tr>
  <tr>
    <td>253</td>
    <td>'no noticeable groove present'<br>'deep medially with distinct groove or wall',</td>
  </tr>
  <tr>
    <td>254</td>
    <td>dorsally positioned on body<br>intermediate condition<br>sunk into body plantarly,</td>
  </tr>
  <tr>
    <td>255</td>
    <td>'relatively small (sliver, oval, no clear insertion<br>for tendo calcanei)'</td>
  </tr>
</table>
'medium (large in one direction, insertion for tendo calcanei present but not robust)'
'relatively large',
256
  oval
  'square / rectangular'
  'uneven walls / trapezoidal'
  'proximally projecting medial wall',
257
  'tucked under ectal facet in lateral view'
  'level with ectal facet in lateral view'
  'projects dorsal to ectal facet',
258
  '(<100)'
  '(100-150)'
  '(>150% longer than wide)',
259
  absent
  present,
260
  'cuboid facet contacts only the ectocuneiform'
  'cuboid facet contacts the ectocuneiform and the mesocuneiform',
261
  wide
  narrow,
262
  '<=105 degrees'
  '>=105 degrees',
263
  'slight concavity'
  'deep concavity, saddle-shaped',
264
  dorsoventral
  '0 / 2'
  intermediate
  '2 / 4'
  mediolateral,
265
  flat
  short
  intermediate
  long,
266
  flat
  shallow
  deep,
267
  'rounded or weak keel'
  'strong keel',
268
  deep
  intermediate
  shallow,
269
  short
  long,
270
  ovoid
  rectangular,
271
<table>
  <tr>
    <td>'medial epicondyle larger'</td>
    <td></td>
  </tr>
  <tr>
    <td>'epicondyles symmetrical'</td>
    <td></td>
  </tr>
  <tr>
    <td>'lateral epicondyle larger',</td>
    <td>272</td>
  </tr>
  <tr>
    <td>medial</td>
    <td></td>
  </tr>
  <tr>
    <td>intermediate</td>
    <td></td>
  </tr>
  <tr>
    <td>lateral,</td>
    <td></td>
  </tr>
  <tr>
    <td>absent</td>
    <td>273</td>
  </tr>
  <tr>
    <td>medioplantar</td>
    <td></td>
  </tr>
  <tr>
    <td>lateroplantar,</td>
    <td></td>
  </tr>
  <tr>
    <td>single</td>
    <td>274</td>
  </tr>
  <tr>
    <td>bifurcate,</td>
    <td></td>
  </tr>
  <tr>
    <td>'smaller or equal in size to peroneal process'</td>
    <td>275</td>
  </tr>
  <tr>
    <td>'larger than peroneal process',</td>
    <td></td>
  </tr>
  <tr>
    <td>present</td>
    <td>276</td>
  </tr>
  <tr>
    <td>absent,</td>
    <td></td>
  </tr>
  <tr>
    <td>'small'</td>
    <td>277</td>
  </tr>
  <tr>
    <td>'hypertrophied',</td>
    <td></td>
  </tr>
  <tr>
    <td>mesaxonic</td>
    <td>278</td>
  </tr>
  <tr>
    <td>paraxonic</td>
    <td></td>
  </tr>
  <tr>
    <td>ectaxonic,</td>
    <td></td>
  </tr>
  <tr>
    <td>'&lt;1.0'</td>
    <td>279</td>
  </tr>
  <tr>
    <td>'0 / 2'</td>
    <td></td>
  </tr>
  <tr>
    <td>'&gt;1.01, &lt;1.5'</td>
    <td></td>
  </tr>
  <tr>
    <td>'2 / 4'</td>
    <td></td>
  </tr>
  <tr>
    <td>'&gt;1.51, &lt;2.0'</td>
    <td></td>
  </tr>
  <tr>
    <td>'4 / 6'</td>
    <td></td>
  </tr>
  <tr>
    <td>'&gt;2.01',</td>
    <td></td>
  </tr>
  <tr>
    <td>'less than 70% the length of MT3'</td>
    <td>280</td>
  </tr>
  <tr>
    <td>'0 / 2'</td>
    <td></td>
  </tr>
  <tr>
    <td>'71-80%'</td>
    <td></td>
  </tr>
  <tr>
    <td>'2 / 4'</td>
    <td></td>
  </tr>
  <tr>
    <td>'81-90%'</td>
    <td></td>
  </tr>
  <tr>
    <td>'4 / 6'</td>
    <td></td>
  </tr>
  <tr>
    <td>'91-100%'</td>
    <td></td>
  </tr>
  <tr>
    <td>'6 / 8'</td>
    <td></td>
  </tr>
  <tr>
    <td>'&gt;100%',</td>
    <td></td>
  </tr>
  <tr>
    <td>'absent or poorly developed'</td>
    <td>281</td>
  </tr>
  <tr>
    <td>'present and distinct',</td>
    <td></td>
  </tr>
  <tr>
    <td>'absent, articular surface of tibial condyles</td>
    <td>282</td>
  </tr>
  <tr>
    <td>perpendicular to longaxis of tibial shaft'</td>
    <td></td>
  </tr>
  <tr>
    <td>'moderate retroflexion present'</td>
    <td></td>
  </tr>
  <tr>
    <td>'proximal articular surface is strongly retroflexed',</td>
    <td></td>
  </tr>
  <tr>
    <td>'absent, small articulation'</td>
    <td>283</td>
  </tr>
  <tr>
    <td>'no fusion, moderate articulation'</td>
    <td></td>
  </tr>
  <tr>
    <td>'no fusion but extensive articulation'</td>
    <td></td>
  </tr>
  <tr>
    <td>fusion,</td>
    <td></td>
  </tr>
  <tr>
    <td>square</td>
    <td>284</td>
  </tr>
</table>
<table>
  <tr>
    <td>285</td>
    <td>intermediate<br>triangular,<br>none<br>'0 / 2'<br>slight<br>'2 / 4'<br>strong,</td>
  </tr>
  <tr>
    <td>286</td>
    <td>flat<br>'0 / 2'<br>'anteriorly convex, posteriorly flat'<br>'2 / 4'<br>'all convex',</td>
  </tr>
  <tr>
    <td>287</td>
    <td>'no compression'<br>'anteroposteriorly compressed',</td>
  </tr>
  <tr>
    <td>288</td>
    <td>'on medial side of malleolus'<br>'0 / 2'<br>'on posterior side of malleolus',</td>
  </tr>
  <tr>
    <td>289</td>
    <td>'small and insignificant'<br>'0 / 2'<br>'present and well-developed',</td>
  </tr>
  <tr>
    <td>290</td>
    <td>spherical<br>semicylindrical<br>cylindrical,</td>
  </tr>
  <tr>
    <td>291</td>
    <td>'restricted to femoral head'<br>'extends onto back of femoral neck',</td>
  </tr>
  <tr>
    <td>292</td>
    <td>'&lt;=75'<br>'0 / 2'<br>'&gt;75, &lt;120'<br>'2 / 4'<br>'=&gt;120',</td>
  </tr>
  <tr>
    <td>293</td>
    <td>'&lt;60'<br>'0 / 2'<br>'60-70'<br>'2 / 4'<br>'&gt;70',</td>
  </tr>
  <tr>
    <td>294</td>
    <td>'bent anteriorly'<br>'not bent anteriorly',</td>
  </tr>
  <tr>
    <td>295</td>
    <td>absent<br>present,</td>
  </tr>
  <tr>
    <td>296</td>
    <td>'hook-shaped'<br>'0 / 2'<br>'rounded',</td>
  </tr>
  <tr>
    <td>297</td>
    <td>absent<br>'0 / 2'<br>present,</td>
  </tr>
  <tr>
    <td>298</td>
    <td>'&gt;125 (long)'<br>'0 / 2'</td>
  </tr>
</table>
'110-125 (moderate)'
'2 / 4'
'<110 (very short)',
299
absent
present,
300
'medial (0-30 degrees)'
'0 / 2'
'posterior (>30 degrees)',
301
'present, large'
'0 / 2'
'present, small and reduced to a thin crest'
'2 / 4'
absent,
302
'proximal to lesser trochanter'
'0 / 2'
'at level of lesser trochanter'
'2 / 4'
'distal to lesser trochanter',
303
absent
present,
304
'100 or higher'
'90-99'
'80-89'
'70-79',
305
'sternal head elongate '
'shortened and triangular in shape',
306
'absent '
'present ',
307
'17 or fewer'
'18'
'0 / 2'
'19'
'2 / 4'
'20'
'4 / 6'
'21 or more',
308
'tuberosity not expressed'
'small tubercle present along margo acetabuli'
'well-developed tuberosity present',
309
'narrow '
'0 / 2'
'markedly flared ',
310
'absent '
'present ',
311
'60-70'
'0 / 2'
'50-59'
'2 / 4'
<table>
  <tbody>
    <tr>
      <td></td>
      <td>'40-49'</td>
    </tr>
    <tr>
      <td></td>
      <td>'4 / 6'</td>
    </tr>
    <tr>
      <td></td>
      <td>'30-39 or &lt;,&rdquo;</td>
    </tr>
    <tr>
      <td>312</td>
      <td>'absent'</td>
    </tr>
    <tr>
      <td></td>
      <td>'present',</td>
    </tr>
    <tr>
      <td>313</td>
      <td>'long tail'</td>
    </tr>
    <tr>
      <td></td>
      <td>reduced</td>
    </tr>
    <tr>
      <td></td>
      <td>absent,</td>
    </tr>
    <tr>
      <td>314</td>
      <td>'wide and biconcave'</td>
    </tr>
    <tr>
      <td></td>
      <td>'anteroposteriorly oriented trough',</td>
    </tr>
    <tr>
      <td>315</td>
      <td>'present'</td>
    </tr>
    <tr>
      <td></td>
      <td>'absent',</td>
    </tr>
    <tr>
      <td>316</td>
      <td>'not confluent'</td>
    </tr>
    <tr>
      <td></td>
      <td>'0 / 2'</td>
    </tr>
    <tr>
      <td></td>
      <td>'partially fused'</td>
    </tr>
    <tr>
      <td></td>
      <td>'2 / 4'</td>
    </tr>
    <tr>
      <td></td>
      <td>'postglenoid process fused to lateral aspect of</td>
    </tr>
    <tr>
      <td>bulla',</td>
      <td></td>
    </tr>
    <tr>
      <td>317</td>
      <td>present</td>
    </tr>
    <tr>
      <td></td>
      <td>absent,</td>
    </tr>
    <tr>
      <td>318</td>
      <td>'posterior to postglenoid process'</td>
    </tr>
    <tr>
      <td></td>
      <td>'0 / 2'</td>
    </tr>
    <tr>
      <td></td>
      <td>'medial to postglenoid process',</td>
    </tr>
    <tr>
      <td>319</td>
      <td>'indistinct or absent'</td>
    </tr>
    <tr>
      <td></td>
      <td>'0 / 2'</td>
    </tr>
    <tr>
      <td></td>
      <td>robust,</td>
    </tr>
    <tr>
      <td>320</td>
      <td>'medially placed'</td>
    </tr>
    <tr>
      <td></td>
      <td>'0 / 2'</td>
    </tr>
    <tr>
      <td></td>
      <td>'laterally placed',</td>
    </tr>
    <tr>
      <td>321</td>
      <td>'medial to space between M1 and M2 or more</td>
    </tr>
    <tr>
      <td>posterior'</td>
      <td>'0 / 2'</td>
    </tr>
    <tr>
      <td></td>
      <td>'medial to M1'</td>
    </tr>
    <tr>
      <td></td>
      <td>'2 / 4'</td>
    </tr>
    <tr>
      <td></td>
      <td>'medial to the space between P4 and M1',</td>
    </tr>
    <tr>
      <td>322</td>
      <td>present</td>
    </tr>
    <tr>
      <td></td>
      <td>'0 / 2'</td>
    </tr>
    <tr>
      <td></td>
      <td>absent,</td>
    </tr>
    <tr>
      <td>323</td>
      <td>absent</td>
    </tr>
    <tr>
      <td></td>
      <td>present,</td>
    </tr>
    <tr>
      <td>324</td>
      <td>'anterior to M3'</td>
    </tr>
    <tr>
      <td></td>
      <td>'0 / 2'</td>
    </tr>
    <tr>
      <td></td>
      <td>'medial to M3'</td>
    </tr>
    <tr>
      <td></td>
      <td>'2 / 4'</td>
    </tr>
    <tr>
      <td></td>
      <td>'posterior to M3',</td>
    </tr>
    <tr>
      <td>325</td>
      <td>'long medial pterygoid plate extending one-third to one half of the distance to the anterior surface of the bulla'</td>
    </tr>
  </tbody>
</table>
its entiredorsoventral extent'
'2 / 4'
'medial pterygoid plate entirely absent, or reduced

to a low rugosity',

326
absent
'0 / 2'
abutting
'2 / 4'
laminar,

327
slight
'very extensive',

328
'lateral side of lateral pterygoid '
'0 / 2, or posterior to terminus of lateral
pterygoid'
'medial to lateral pterygoid ',

329
narrow
broad,

330
broad
peaked,

331
'absent or minimal'
'present, extensive',

332
absent
'present and formed by the AAC'
'present and formed by the tympanic cavity'
'1 / 2',

333
tubular
'0 / 2'
'not tubular',

334
absent
'present, small, and in the posterior root of the
'present, large, and above the external auditory
zygomatic arch'

meatus'

335
'if present is poorly developed '
'present and enlarged ',

336
'absent '
'present ',

337
'ventral to m. tensor tympani'
'dorsal to m. tensor tympani'
'm. tensor tympani absent',

338
'LN(PC diameter / SC diameter) <0'
'0 / 2'
'LN(PC diameter / SC diameter) >0.01, <1.0'
'2 / 4'
'LN(PC diameter / SC diameter) >1.01, <2.0'
'4 / 6'
'LN(PC diameter / SC diameter) >2.01, <10'
'6 / 8'
'LN(PC diameter / SC diameter) >10',
339
medial
'midline of bulla'
lateral,
340
'posterior to fenestra cochleae'
'ventral to fenestra cochleae'
'anterior to fenestra cochleae',
341
dorsal
ventral,
342
'runs across ventral lip of fenestra cochleae,
shielding it from ventral view'
'does not shield fenestra cochleae',
343
'promontorial canal present and fully enclosed in
its course through the middle ear'
'promontorial canal present but short and continuous
anteriorly with trough / groove'
'promontorial canal absent, trough or groove present
across entire promontorium',
344
absent
present,
345
absent
'tympanic cavity'
'present, forms lateral wall of AAC pneumatized from
epitympanic recess',
346
absent
present,
347
'<1.9'
'0 / 2'
'2.0-2.49'
'2 / 4'
'2.5-2.99'
'4 / 6'
'>3',
348
petrosal
entotympanic,
349
absent
present,
350
'linea semicircularis or partial anular bridge
formed on anentotympanic bulla'
'linea semicircularis formed on a petrosal bulla'
'complete annular bridge',
351
intrabullar
extrabullar,
352
<table>
  <tbody>
    <tr>
      <td></td>
      <td>patent</td>
    </tr>
    <tr>
      <td></td>
      <td>closed,</td>
    </tr>
    <tr>
      <td>353</td>
      <td>absent</td>
    </tr>
    <tr>
      <td></td>
      <td>'present, moderately inflated'</td>
    </tr>
    <tr>
      <td></td>
      <td>'present, greatly inflated',</td>
    </tr>
    <tr>
      <td>354</td>
      <td>absent</td>
    </tr>
    <tr>
      <td></td>
      <td>'0 / 2'</td>
    </tr>
    <tr>
      <td></td>
      <td>'present, small'</td>
    </tr>
    <tr>
      <td></td>
      <td>'2 / 4'</td>
    </tr>
    <tr>
      <td></td>
      <td>'present, well-developed',</td>
    </tr>
    <tr>
      <td>355</td>
      <td>'present'</td>
    </tr>
    <tr>
      <td></td>
      <td>'0 / 2'</td>
    </tr>
    <tr>
      <td></td>
      <td>'absent',</td>
    </tr>
    <tr>
      <td>356</td>
      <td>'temporal lines converge on frontal'</td>
    </tr>
    <tr>
      <td></td>
      <td>'temporal lines converge on the parietals, forming</td>
    </tr>
    <tr>
      <td>short sagittalcrest'</td>
      <td>'temporal lines are not confluent, no sagittal crest</td>
    </tr>
    <tr>
      <td>present'</td>
      <td>'sagittal crest is extensive, but temporal lines</td>
    </tr>
    <tr>
      <td>converge onparietals that extend far rostrally between frontal bones'</td>
      <td>'0 / 2',</td>
    </tr>
    <tr>
      <td>357</td>
      <td>absent</td>
    </tr>
    <tr>
      <td></td>
      <td>'0 / 2'</td>
    </tr>
    <tr>
      <td></td>
      <td>present,</td>
    </tr>
    <tr>
      <td>358</td>
      <td>present</td>
    </tr>
    <tr>
      <td></td>
      <td>'absent due to intervening lacrimal'</td>
    </tr>
    <tr>
      <td></td>
      <td>'absent due to intervening premaxilla',</td>
    </tr>
    <tr>
      <td>359</td>
      <td>'&gt;89'</td>
    </tr>
    <tr>
      <td></td>
      <td>'0 / 2'</td>
    </tr>
    <tr>
      <td></td>
      <td>'90-104'</td>
    </tr>
    <tr>
      <td></td>
      <td>'2 / 4'</td>
    </tr>
    <tr>
      <td></td>
      <td>'105-119'</td>
    </tr>
    <tr>
      <td></td>
      <td>'4 / 6'</td>
    </tr>
    <tr>
      <td></td>
      <td>'120-134'</td>
    </tr>
    <tr>
      <td></td>
      <td>'6 / 8'</td>
    </tr>
    <tr>
      <td></td>
      <td>'&gt;135',</td>
    </tr>
    <tr>
      <td>360</td>
      <td>'no anteromedial expansion'</td>
    </tr>
    <tr>
      <td></td>
      <td>'anterior and medial expansion',</td>
    </tr>
    <tr>
      <td>361</td>
      <td>'extensive transverse lamina and ethmoturbinal</td>
    </tr>
    <tr>
      <td>recess present'</td>
      <td>'extensive transverse lamina and ethmoturbinal</td>
    </tr>
    <tr>
      <td>recess absent',</td>
      <td></td>
    </tr>
    <tr>
      <td>362</td>
      <td>absent</td>
    </tr>
    <tr>
      <td></td>
      <td>'present below CN I',</td>
    </tr>
    <tr>
      <td>363</td>
      <td>small</td>
    </tr>
    <tr>
      <td></td>
      <td>large</td>
    </tr>
    <tr>
      <td></td>
      <td>hypertrophied,</td>
    </tr>
    <tr>
      <td>364</td>
      <td>'confluent with the superior orbital fissure'</td>
    </tr>
    <tr>
      <td></td>
      <td>'present',</td>
    </tr>
  </tbody>
</table>
365
'surrounded by lacrimal bone '
'foramen lies on lacrimal / maxillary suture '
'foramen lies on the lacrimal / maxillary suture but
is primarily surrounded by maxilla ',
366
'on rim, or outside of, orbit '
'0 / 2'
'inside the orbit ',
367
'present '
'0 / 2'
'absent ',
368
'contact present '
'no contact, separated by a large fronto-maxillary
contact (and insome taxa, a small os planum of the ethmoid) '
'no contact, separated by a large os planum',
369
'no complete ethmomaxillary fissure present'
'0 / 2'
'complete ethmomaxillary fissure present',
370
'anterior hiatus variably present'
'0 / 2'
'posterior hiatus at least variably present'
'2 / 4'
'no hiatuses present',
371
present
'0 / 2'
absent,
372
'within palatine'
'0 / 2'
'on ethmopalatine suture',
373
'above P2'
'0 / 2'
'above P3'
'2 / 4'
'above P4'
'4 / 6'
'above M1',
374
'no postorbital bar '
'slender and delicate '
'1 / 3'
'wide and robust '
'contributes to postorbital septum ',
375
'partial frontal process but no postorbital bar '
'slender and delicate '
'1 / 3'
'wide and robust '
'incorporated into postorbital septum ',
376
slender
'0 / 2'
'dorsoventrally deep',
377
<table>
  <tbody>
    <tr>
      <td></td>
      <td>'absent'</td>
    </tr>
    <tr>
      <td></td>
      <td>'0 / 2'</td>
    </tr>
    <tr>
      <td></td>
      <td>'present, small'</td>
    </tr>
    <tr>
      <td></td>
      <td>'2 / 4'</td>
    </tr>
    <tr>
      <td></td>
      <td>'present, large',</td>
    </tr>
    <tr>
      <td>378</td>
      <td>'long snout'</td>
    </tr>
    <tr>
      <td></td>
      <td>'short snout',</td>
    </tr>
    <tr>
      <td>379</td>
      <td>deep</td>
    </tr>
    <tr>
      <td></td>
      <td>shallow,</td>
    </tr>
    <tr>
      <td>380</td>
      <td>straight</td>
    </tr>
    <tr>
      <td></td>
      <td>'0 / 2'</td>
    </tr>
    <tr>
      <td></td>
      <td>dished,</td>
    </tr>
    <tr>
      <td>381</td>
      <td>'foramina sit behind first incisors'</td>
    </tr>
    <tr>
      <td></td>
      <td>'0 / 2'</td>
    </tr>
    <tr>
      <td></td>
      <td>'foramina intrude between first incisors',</td>
    </tr>
    <tr>
      <td>382</td>
      <td>'no extension of premaxilla beyond anterior'</td>
    </tr>
    <tr>
      <td>incisors'</td>
      <td>'mild premaxillary projection'</td>
    </tr>
    <tr>
      <td>rostrum',</td>
      <td>'considerable projection, forming a tubular anterior'</td>
    </tr>
    <tr>
      <td>383</td>
      <td>narrow</td>
    </tr>
    <tr>
      <td></td>
      <td>broad,</td>
    </tr>
    <tr>
      <td>384</td>
      <td>'absent'</td>
    </tr>
    <tr>
      <td></td>
      <td>'present',</td>
    </tr>
    <tr>
      <td>385</td>
      <td>'lateral cleft between medial and lateral nasal'</td>
    </tr>
    <tr>
      <td>processes'</td>
      <td>'lateral cleft fused and rhinarium covered with dry'</td>
    </tr>
    <tr>
      <td>hairy skin',</td>
      <td></td>
    </tr>
    <tr>
      <td>386</td>
      <td>'absent'</td>
    </tr>
    <tr>
      <td></td>
      <td>'present',</td>
    </tr>
    <tr>
      <td>387</td>
      <td>'present'</td>
    </tr>
    <tr>
      <td></td>
      <td>'absent',</td>
    </tr>
    <tr>
      <td>388</td>
      <td>'choriovitelline placenta'</td>
    </tr>
    <tr>
      <td></td>
      <td>'no choriovitelline placenta',</td>
    </tr>
    <tr>
      <td>389</td>
      <td>epitheliochorial</td>
    </tr>
    <tr>
      <td></td>
      <td>'diffuse, hemochorial',</td>
    </tr>
    <tr>
      <td>390</td>
      <td>invasive</td>
    </tr>
    <tr>
      <td></td>
      <td>'non-invasive',</td>
    </tr>
    <tr>
      <td>391</td>
      <td>'primordial cavity absent'</td>
    </tr>
    <tr>
      <td></td>
      <td>'primordial cavity present',</td>
    </tr>
    <tr>
      <td>392</td>
      <td>absent</td>
    </tr>
    <tr>
      <td></td>
      <td>present,</td>
    </tr>
    <tr>
      <td>393</td>
      <td>'large, vesicular'</td>
    </tr>
    <tr>
      <td></td>
      <td>rudimentary,</td>
    </tr>
    <tr>
      <td>394</td>
      <td></td>
    </tr>
  </tbody>
</table>
absent
present,

;

### MATRIX

'Tupaia glis' 0000404441 0000000003 2204042022 0202612002
2200120150 0000008066 4300002004 2402442202 1040007441 0020210000 00200240-
0 2012000211 2102-00040 2010000000 00200100-3 00-0020002 4603031004
0100000000 0140010300 4302420202 1344010001 0102204242 4000260808
4021010100 0130000001 0001110000 1000000010 0101000160 0102230010
0311001{13}00 040100{246}200 0000000010 2004000-10 0011002100 0100005110
0002011000 0000000004 0011104010 001?000000 0001

'Absarokius sp.' 10004??2?? ??00?00002 2202041022 0210212004 -
200120250 4432000802 4200122044 2212352102 3440025262 1122210021 ???? ???????
?????????12 ?1?2220041 0000000000 4126020204 -0-0202202 0200040200
3210000??0 ??????????? ?002020220 024??????? ?220020242 2026246244
2020010102 1101000100 1011110??? ??????????? ??????????? ??202212??
?????????? ??????????? ??????????? ??????????? ??????????? ???? ???????
?????????? ??????????? ??????????? ??????????? ?????

'Abuqatrania basiodontos' ??0???????? ???? ??????? 22??????2 2????12004

100000?260 ?2?0002022 2100222110 2202222000 1202045120 1022210200
?????????? ??????????? ??????????? ??????????? ??????????? ???? ???????
?????????? ???????????00 02???????? ??????????? ??????????? ???? ???????
?????????? ??????????? ??????????? ??????????? ??????????? ???? ???????
?????????? ??????????? ??????????? ??????????? ??????????? ???? ???????
?????????? ??????????? ??????????? ??????????? ?????

'Adapis parisiensis' 1000200000 1402?00223 0004440022 0004414204 -
200044404 1020226668 0-00242405 24200-2222 40500000-2 2202210040 ?204040120
4214004001 2104000140 0020121000 2322020200 00-0000002 0404000031
0420402422 01???20420 3302320001 024602?04? 1530321104 300743{13}308
4011200032 0101011101 1111120??? 0112221210 0110010??? 0102440210
0200001200 1112?????? ??0030002 2013420010 0020???020 0010002?11
011403106? ?0011001?? ??33322100 001??????? ?????

'Aegyptopithecus zeuxis' 1000202000 1102200003 24-0-40022 2216814214 -
411224231 1300100208 0-01140020 11320-2000 32020330-1 2022210020 1?????????
?112012613 ---0042042 0010201100 41210220-0 00-1000000 0{024}02100342
2101402204 12???20420 2322020212 1048?????? 0642404744 4003442????
4011010020 ?1220??201 1012?????? 0001110210 2020011??? ???? ???????0
??01000002 1402??????? ????0000021 110320-?11 0020???812 1101114?0-
1110?4208? ?10?112??? ??64402002 001??????? ?????

'Afradapis longicristatus' 1001?00100 1102?00203 23---40022 2216814004 -
204030422 2000208668 0-00222225 24300-3222 42500020-2 2002210022 1??1040010
4?12?12613 ---0120042 0020001000 1120000200 00-4001001 4404001031
0314?02224 ???? ??????? ???? ??????? ???? ??????? ?640044024 4108420???
?????????? ???? ??????? ???? ??????? ???? ??????? ???? ??????? ???? ???????
?????????? ???? ??????? ???? ??????? ???? ??????? ???? ??????? ???? ???????
?????????? ???? ??????? ??4??????? ?????

'Aframonius dieides' 1?0??0000? 1??2?00003 2201042112 0015713114 -
203132421 0100105358 0-002235-6 24000-3232 42500010-2 4000210031 ???? ???????
?????????12 2102020040 2020001000 4120020200 00-1022001 2402001011
0422401201 01???????? ???? ??????? ???? ??????? ???? ??????? ???? ???????
?????????? ???? ??????? ???? ??????? ???? ??????? ???? ??????? ???? ???????
?????????? ???? ??????? ???? ??????? 2010??????? ???? ??????? ???? ???????
?????????? ??1??0???? ??41??2??? ?????

'Algeripithecus minutus' ???? ??????? ???? ??????? 22???40022 0226610--4 -
004042420 2040228851 0-002046-6 23220-2332 4-600140-3 2002210010 ???? ???????
?????????2 21??020242 0021020100 002???0200 00-2001000 0402100432
2201?0???0 ???? ??????? ???? ??????? ???? ??????? ???? ??????? ???? ???????
?????????? ???? ??????? ???? ??????? ???? ??????? ???? ??????? ?????
3023202203 01???20410 3122022212 1212??1??? ?320124644 3003066808
4000010202 1220000200 10011100?0 ???? ??????? ????1??? 0120220220
0411011012 3200???00? ???? ??????? ??10?????? ??2??????02 ??011???0?
111??02?4? ?10??12??? ??3?4?2?0? ???? ??????? ?????
'Arapahovius gazini' 1102804441 1440?20010 2202042222 0242613010
2202240230 2241204122 4200122634 2342352122 2350002261 2102310024
???30420?? ???? ????2?? 21?2220040 0020000000 4120020204 -0-0304222
0401040020 2320300??? ???? ??????? ???? ??????? ???? ??????? 10400???42
002?0??242 4020110202 1101000000 00001102?0 1113320210 201001????
?????????? ???? ??????? ???? ??????? ???? ??????? ???? ???????
?????????? ???? ??????? ???? ??????? ???? ??????? ?????
'Arctocebus calabarensis' 1010414241 1200011034 2208040022 2206610--4 -
204020110 2020226068 0-002222-6 24410-2222 4-600030-2 2002210000 0002022000
2113102211 2122020042 0020100200 4320020100 00-2000002 0403000142
0101200200 0100502401 4420420200 3216?021?1 1612462044 4108410524 4020--1--
0 0022022011 3100000?0? 0114320201 111001?208 0102440200 1010000300
3003118000 6120020012 210220-110 001011?-00 0?2020?00- 11202200?1
0010102200 2233111011 21010000?? ?????
'Arsinoea kallimos' 1001002000 1402?10003 2204040222 2024412004 -
200020240 4220020224 2300242002 0202252020 20210240-2 0022210020 ???? ???????
?????????? ???? ??????? ???? ??????? ????22000-0 00-0000002 0202200242
0000000??2 ???? ??????? ???? ??????? ???? ??????? ???? ???????
?????????? ???? ??????? ???? ??????? ???? ??????? ???? ???????
?????????? ???? ??????? ???? ??????? ???? ??????? ???? ???????
??????2?6? ???? ??????? ???? ??????? ?????
'Asiadapis cambayensis' ???? ??????? ???? ??????? 22???40022
0226811004 2202020220 0020118464 3100212244 2222152022 2440005162
3110210101 ???? ??????? ???? ??????? ????020040 0000000000 2024020202 00-
0001000 0202040010 0220200??0 ???? ??????? ???? ??????? ???? ??????? ?240020242
4046400806 202?000122 1102000000 11101?0??? ???? ??????? ???? ???????
?????????00 1?00022000 240??????? ???? ??????? ???? ??????? ?????
?????????? ???? ??????? ???? ??????? ???? ??????? ?????
'Azibius trerki' ???? ??????? ???? ??????? ???? ???????2 ?0?6?10--2
0004022220 2020225852 0-002246-6 23200-2232 4-600220-3 2002210010
?????????? ???? ???????12 ????020040 0022020200 6124020200 00-0020002
0202000202 220020??? ???? ??????? ???? ??????? ???? ??????? ?22?0???40
202?4????? ???? ??????? ???? ??????? ???? ??????? ???? ???????
?????????? ???? ??????? ???? ??????? ???? ??????? ???? ???????
?????????? ???? ??????? ???? ??????? ???? ??????? ?????
'Bahinia pondaungensis' 1?00000000 1204?10003 22040?0022 222?810--4 -
200020222 0000014644 4102200044 2202?????? ????0?422 2022210010 ???? ???????
?010012212 0102000040 002000?000 ?1220220-4 00-00000000 0402031242
0?????0??4 ???? ??????? ???? ??????? ???? ??????? ???? ???????
?????????? ???? ??????? ???? ??????? ???? ??????? ???? ???????
?????????? ???? ??????? ???? ??????? ???? ??????? ???? ???????
?????????? ???? ??????? ???? ??????? ???? ??????? ?????
'Bahinia banyueae' ???? ??????? ???? ??????? ????40022 ?02????????
?????????? ???? ??????? 43002?0?? ?222422022 044000???? ????0020
?????????? ???? ??????? 01?02000?0 0020000000 40?0220-4 -0-0000000
05?2040042 04000????? ???? ??????? ???? ??????? ???? ??????? ???? ???????
?????????? ???? ??????? ???? ??????? ???? ??????? ???? ???????
?????????? ???? ??????? ???? ??????? ???? ??????? ?????
'Biretia fayumensis' ?0?????0?? ???? ??????? 22???40022 202??12004 -
200030240 ?100005252 2101221123 2202122012 22320240-3 0022210011 ???? ???????
?????????? ????042042 0001000200 212?0?0100 00-0004002 0002200240
2?????0??? ???? ??????? ???? ??????? ???? ??????? ???? ???????
?????????? ???? ??????? ???? ??????? ???? ??????? ?????
?????????? ???? ??????? ???? ??????? ?????
?????????? ???? ??????? ?????
'Biretia megalopsis' 1?0?????0?? ??????0??? 2204040022 2026813104 -
20013?140 2200007?53 3101222222 2212121022 1222035123 0022210020 ??????????
???0???12 ???????42 0001001200 2122020200 00-0034002 0002200241
320020???2 ??????????? ??????????? ??????????? ??????????? ???????????
?????????? ??????????? ??????????? ??????????? ??????????? ???????????
?????????? ??????????? ??????????? ??????????? ??????????? ???????????
?????????? ??????????? ?11??????? ??????????? ?????

'Branisella boliviana' 1?0?????000 1?02110?03 2204042220 -226404224 -
4?0224220 1200000006 0-02040404 22020-2032 20500260-0 2020210020 ??????????
???1??12 ???2240020 2000200000 01100200-0 00-0030002 0002100244
2000022??4 ??????????? ??????????? ??????????? ??????????? ???????????
?????????? ??????????? ??????????? ??????????? ??????????? ???????????
?????????? ??????????? ??????????? ??????????? ??????????? ???????????
?????????? ??????????? ?11??????? ??????????? ?????

'Bugtilemur mathesoni' ??0??????? ?????2020? ?????40022 ?22??14222

020444?404 ?020008?62 4400240024 2402252002 222010???? ????0032
?????????? ??????????? 01??0000?22 ??0022??0? ????0?0104 ---0300001
00?4040040 0?????????? ??????????? ??????????? ??????????? ???????????
?????????? ??????????? ??????????? ??????????? ??????????? ???????????
?????????? ??????????? ??????????? ??????????? ??????????? ?????

'Caenopithecus lemuroides' ??0??????? ?????????? 2200040022 2226612004 -
202020411 2200108268 1000242225 24200-2122 34501220-2 2000310042 ??????????
?????????? ????000?40 0020001000 2120020200 00-2002002 4404021022
0220402??2 ??????????? ??????????? ????4?040??4 ??08440606
2000100112 0012011011 1100220??? ??????????? ??????????? ???????????
?????????? ??????????? ??????????? 01?1??????? ??????????? ???????????
?????????? ??0??????? ??41?22??? ??????????? ?????

'Cantius abditus' 1000000220 1202?00?03 0004040022 0224612002

1201220222 0000106408 3300213544 2432253222 3450010263 3202410021
??????????? ???200?001 2102240040 1020101010 4122021200 00-0442002
1402000030 0210200022 01???0?42? ?302400?40 004??????? ?340221142
?045?14526 0000100100 0201000100 0100110??? ???330???0??? ??????????
???0440??0 0????0???? 0211?????? ??000?000? ??2?????1? 00??????020
000000??1? ?1???????? ?01??????? ??3??????? ?1??????? ?????

'Cantius ralstoni' ????0?????? ????0?????? 00???40022 0224613004 -
201220220 0000005228 4300203544 2222452232 2440011263 2204410022 ??????????
?????????? ????240040 0020001000 4023022204 -0-0442002 0402040030
221020???2 ????0?????? ????0?????? ????0?????? ?422203?? ?045405?26
0000110102 0102000001 110011?101 ????0?????? ????0?????? ????0???????
?????????? ????0?????? ????0?????? ????0?????? ????0?????? ????0??????
?????????? ????0?????? ????0?????? ????0?????? ????0?????? ?????

'Carpolestes simpsoni' 1?02804441 0040020010 24-0------ ----

-10--4 0004002420 22{13}2008850 40002046-6 2312442202 2-60000261 4002410020
10400042-1 420------12 2102042240 2002020212 2026000202 00-0204222
0402030000 0220400024 01???00400 0302020020 0444???0?1 ?202004802
4004062806 40210000200 0230000011 1101100000 0002001210 02?0000?26
0200040020 1421000000 0403???100 0000020020 2003?20000 ?021???01
?120000?0? ??00?1000? ?0010021?? ??30???0??0 201?????? ?????

'Catopithecus browni' 1000101000 1002200003 24-0-20022 2205714222

1402234221 1100003318 2101140002 22010-2002 20210150-1 2020410020
1221040000 2012010613 ---0220040 0010200000 41220210-2 00-0000000
0302120242 2200200202 02???????0 3322020202 102??????? ?441322444
4004340??? ????0?????? ????0?????? ????0?????? 000111?210 202001????
???????20 02{13}1000002 04??????? ??00000022 ?21?????1? 0?20????02
110?????? 111??12?6? ?10??12??? ??3440210? 001?????? ?????

'Cheirogaleus major' 1010414241 1200011034 2207141020 -116611004 -
20432{13}203 1140004236 0-002403-6 24320-2032 3-600050-3 3222210022
0241042010 0214100212 2122000042 0000100000 10200100-4 -0-0030002
0103040042 2300200000 0155320440 0101420241 00431?1031 1220324241
3028402224 2121210022 0112001000 1100130101 0112321210 2210010226
'Lemur catta' 1010414241 1200011034 2207040022 2202212204 -
202100220 0200008668 21022420-6 24000-4102 4-600140-1 4022210020 0030030000
2014000211 2103-00040 0000000000 40223100-3 20-0000002 0004230041
2400200000 0166310420 2012410131 1242121021 16203401{13}4 4037426328
3000200022 0111000000 2100221101 0112321210 2210010225 0101440200
0341000100 031000{246}220 4000010002 2001240010 0020000020 101000?011
0103122030 0001100004 0042204010 1010000000 000?

'Leontopithecus rosalia' 1001200200 1004000003 2208042000 -224704224 -
420224222 2200002807 0-020402-6 22020-2032 4-60008--- ------0020 1224020000
2012012212 0024040022 0000201000 41120000-4 -0-0000000 0104040013 0------
2424 1206202201 4420220220 12460110?1 1210204844 4002060826 4111010112
1121001101 101211200? 1001111210 001001?260 0100020020 1401102000
2203004100 2000000202 421-20-110 002000??02 110111?00- 1110222060
1101112204 2024402100 000?111??? ?????

'Lepilemur mustelinus' 1010414241 1200011134 2208042022

2214814004 -304411413 2030108068 0-00222224 24220-2122 22430040-2
3002210020 ----------- --241?0211 2114120042 0010100000 10220100-3 00-
0020000 0304041014 2300201010 0124420410 0301400240 0040111031

142013{13}143 3027436236 2020110120 0121001000 310212210? 0012221210

221001?224 0202430210 0231001300 0110007220 4000020012 2000120010

00200?0310 101000?011 01032{12}001? 0010201121 0053202??0 ?00?00????? ?????

'Leptadapis magnus' 10002?0000 1402?10203 0004442222 0005814002

0200244404 0000106868 0-002424-6 24300-2222 40400000-2 2202210040
1?0??401?? 4014014001 2102040040 0120221000 2324020200 00-0120002
0304000042 0422402?22 ?1???20420 0322220340 024??????? 1{35}30220114

40464{13}0506 2000100032 1112011011 2100120??? 0112221210 2210010???

010????0??? ???? ??????? ????0040002 00134??010 0020??????20

00?000??11 011403104? ?0011001?? ??3332??00 201??????? ?????

'Leptadapis priscus' ????0000?? ????0000?? ????0000??2 0????14204 -

200044424 00?0206668 0-00242424 22200-2222 42400200-2 4002210000 ????0000???

?????????? ????0000?? ????0000?? ????0000?? ????0000?? ????0000???

?????????24 ????0000?? ????0000?? ????0000?? ????0000?? ????0000???

?????????? ????0000?? ????0000?? ????0000?? ????0000?? ????0000???

?????????? ????0000?? ????0000?? ????0000?? ????0000?? ????0000???

?????????? ????0000?? ????0000?? ????0000?? ????0000?? ????0000???

'Leptadapis ruetimeyeri' ????0000?? ????000??0?? 000?410122 2026614202

0200244424 20?0206668 1000243525 24200-2222 22400220-2 4002210020

?????????? ????0000?? 21?2020040 0022021200 0324020200 10-1030001

1402000001 242220??24 ????0000?? ????0000?? ????0000?? ????0000???

?????????? ????0000?? ????0000?? ????0000?? ????0000?? ????0000???

?????????? ????0000?? ????0000?? ????0000?? ????0000?? ????0000???

?????????? ????0000?? ????0000?? ????0000?? ????0000?? ????0000???

'Loris tardigradus' 1010414241 1100011034 2208041022 2204714214 -

300130121 2110006044 0-022321-6 24120-2212 3-600030-1 2002210010 0100002000

0214200211 2124241040 2000100100 3120020200 1310002001 02020001{13}3

0212100200 0100502201 4200420201 24051021?1 1522333144 4107410522 2120--1--

2 1022022010 3100000001 0114320201 111001?208 0102440220 1111000400

3203118101 6120040022 100320-210 0010110-00 112020?00- 1110220041

0110202221 2143222012 12010000?? ???1

'Loveina zephryi' 10004??3?? ??0??????? 22???2?122 0202812000

2202200222 2431104441 4200222034 2432432202 2350020231 2???210020

?????????? ????0000?? ????0000??0 ?0200?1000 412?020204 -0-0102222

0400041000 0220400??? ????0000?? ????0000?? ????0000?? ????0000???

?????????? ????0000?? ????0000?? ????0000?? ????0000?? ????0000???

?????????? ????0000?? ????0000?? ????0000?? ????0000?? ????0000???

?????????? ????0000?? ????0000?? ????0000?? ????0000?? ????0000???

'Macrotarsius montanus' 1?006??2?? ??2??????? 22??????02

02???14220 1402240222 0401203026 4320142022 2422442202 2220020422

4002210022 ????0000?? ????002?? ????0000?? ????0000?? ????0000???

?????????? ????0000?? ????0000?? ????0000?? ????0000?? ????0000???

?????????? ????0000?? ????0000?? ????0000?? ????0000?? ????0000???
```
'Oligopithecus savagei'              ??0???????   ????10003  24-0-20022  2206814222
240002??222  2200002208  2100040002  22020-2001  004100???   ???????0020
???0??????   ???????????   ????0420??   ?0002??200  4???020-2  00-?000000
00??????44  0??????????4   ???????????   ???????????   ???????????   ???????????
??????????   ???????????   ???????????   ???????????   ???????????   ???????????
??????????   ???????????   ???????????   ???????????   ???????????   ???????????
??????????   ???????????   ???????????   ???????????   ???????????   ???????????
'Omomys sp.'                      1102703221  0420?00002  2202040022  0246812004 -
202240220  0020205444  4300220035  2422222002  2350012221  2202210020  ?2??????0??
?210???212  2102220040  0010000000  0124122202  00-0002212  0302220040
0420100??0   ???????????   ???????????   ???????????   ?310030132  3024065046
4021110112  1101000000  111121120?   ???????????   ???????????   ??1022?0?0
004??0?4?0   0110???10?   0?????????   ??????????0  1?22???000  0000004?1?
?01??22???   ??2??202???   ????1??000  0??????????   ?????
'Otolemur crassicaudatus'           1010414241  1200011034  2208042022  2206812004 -
200244404  1220008668  0-022011-6  24020-2112  2-600160-2  2012210010  0010002000
0214200211  2126210041  0102020201  6323010100  1421000002  0002000204
2200100012  0144400440  0012420220  01421?1041  1430051141  2027446062
2020100022  0112101200  210023021?  0114321211  2210010216  1202300222
1031120300  1110005100  6000010012  300120-110  0020110-00  002020?00-
1120220051  0110101201  2111200010  00000000??   ?????
'Pan troglodytes'                   1002400000  1404200203  24-0-41102  2226814224 -
422244221  0400002128  0-02040044  00020-2000  24221240-1  2022210223  1242040200
20120??2613  ---0040042  0000001000  41100020-0  00-2000000  0004000012
020220220?  1206402401  4412420240  1048202030  1642403724  400325{13}808
4121000120  1121000011  2011210001  0002110210  0120010043  0100020220
0401000012  4-03100020  6020041-22  020420-010  0000001812  110111?00-
1120222080  100{01}112204  2254402100  000011111?  1110
'Parapithecus fraasi'               2-0--?000?   ????2?00003  2204042242  2004612004 -
000002040  0400004628  0-02242220  02220-0200  22020440-0  0022210220  ??????????
????????2??   ???????????   ???????????   ???????????   ???????????   ???????????
?????????0?   ?1?????????   ???????????   ???????????   ???????????   ???????????
??????????   ???????????   ???????????   ???????????   ???????????   ???????????
??????????   ???????????   ???????????   ???????????   ???????????   ???????????
??????????   ???????????   ???????????   ???????????   ???????????   ???????????
'Parapithecus grangeri'            3-0------- ---2?00003  2204042142  2205712013
?0000040?0   1400001408  0-01141121  02120-1100  12120440-0  0022210210
1?????????   ?2121?0211  1002040042  0002001000  4123001220  00-0044002
0303100232  2020102?23   ???????04?0  2122022222  122???????   ?3200???42
?0020?????   ???????????   ???????????   ???????????   ???????????   ???????????
??????????   ???????????   ???????????   ????0020002  221204121?  0020???802
110110???-  1?10?02?2?   ?10??12???   ??34404100  000???????   ?????
'Perodicticus potto'                1010414241  1000011134  2208040031  1203311004 -
202020110  0000018066  0-012211-6  00020-2120  2-600280-3  1122210020  0011030000
4114100212  2146010032  0000000000  4120010102  1121000000  0003020314
2100010002  0102600420  1001320200  3346102141  1422333{35}44  4107410706  4010--
1--2  1022022011  310000000?  0114320201  1210010204  0102440010  1011000400
2203107000  6010010012  2101220110  002011?-01  1?2020300?  1113220051
0010201201  12{35}3221010  00010000??   ????1
'Phenacopithecus xueshii'           ????0????   ???????????  2202240022  220??11004 -
20012??220   ?200104?52  42002?0224  2422222002  024000??221  ?222210000  ??????????
??????????   ???????????   ???????????   ????0??104  -0-?00????   ?????????30
0?00??????   ???????????   ???????????   ???????????   ???????????   ???????????
??????????   ???????????   ???????????   ???????????   ???????????   ???????????
??????????   ???????????   ???????????   ???????????   ???????????   ???????????
??????????   ???????????   ???????????   ???????????   ???????????   ???????????
'Plesiadapis tricuspidens'           2?0--?4441 ?04-------- 24-0?42042  0224810--4 -
200042220  0000002208  4400204622  2422254222  2220020461  4102410000  10400242-1
201???--11  2102140140  0002021000  2122020202  00-0304002  0402020000
2220400004  00?0?00420  0302022222  1248?110?1  04?2004804  4006410808
2010000200  0230000011  000110000?  0000000010  0100101?00  0000000020
```
```
'Siamopithecus eocaenus'              ??0?????0??  ???4?00003 22???40022 2204812004 -
200224220 0200000208 0-02240044 02020-2030 24400200-3 0002210020 ???? ???????
???2?????12  ???020040 0000000000 41220202-2 00-4000000 0000020202
4202220?24  ???? ???????  ???? ???????  ???? ???????  ???? ???????
??????????  ???? ???????  ???? ???????  ???? ???????  ???? ???????
??????????  ???? ???????  ???? ???????  ???? ???????  ???? ???????
?????????8?  ?0?????????  ??24????10?  ???? ???????  ?????
'Steinius spp.'                   11006??3??  ????2??????  02???40022 0244812002
1203240220 0000204242 4300222235 2332353002 2350003262 2212310020
??????????  ???? ????1??  ????220040 2020000000 4124020104 -0-0302222
0400040000 0210200???  ???? ???????  ???? ???????  ???? ??????? ??40030{13}??
??35155???  ???? ???????  ???? ???????  ???? ???????  ???? ???????
??????????  ???? ???????  ???? ???????  ???? ???????  ???? ???????
??????????  ???? ???????  ???? ???????  ???? ???????  ???? ???????
??????????  ???? ???????  ???? ???????  ???? ???????  ?????
'Taqah propliopithecid'            ??0???????  ???2100??  24-0-40022 2226814224 -
402244240 2400000408 2102240402 02020-2000 22220240-3 2022210020 ???? ???????
??????????  ????040040 0001201200 41??2??20-2 00-4000000 0102220242
2????????4  ???? ???????  ???? ???????  ???? ???????  ???? ???????
??????????  ???? ???????  ???? ???????  ???? ???????  ???? ???????
??????????  ???? ???????  ???? ???????  ???? ???????  ???? ???????
??????????  ???? ???????  ???? ???????  ???? ???????  ?????
'Tarsius bancanus'                 2?0--??3111 ?002000200 2202440020 -220211004 -
202220220 4241213114 4100200025 2432422002 2250022421 2102210010 14441040-0
2010012212 3102200042 0020000000 4120010104 -0-0001000 0302040140
1400400200 1166520420 0002420240 0240011021 11000202{13}0 2004061080
2021200032 0100000201 1101231210 1113321201 2020011225 0230221022
1130100400 0010003000 6001020220 2002241001 1102000622 110110600-
1000022010 1121102211 0?13400100 0000111111 1110
'Tarsius sirindhornae'              2?0--??2??  ???? ??????? 22?????0??2
02?2411004 -20002022? 42412040{13}5 4100200024 2442422002 224002242?
2102210020  ???? ???????  ???? ???????  ????200042 0020200000 01200?0204 -0-
?002222 060?????4?  0????????0  ???? ???????  ???? ???????  ???? ???????
??????????  ???? ???????  ???? ???????  ???? ???????  ???? ???????
??????????  ???? ???????  ???? ???????  ???? ???????  ???? ???????
??????????  ???? ???????  ???? ???????  ???? ???????  ?????
'Teilhardina americana'            11016??4??  ??10?00002 1202041022
0221{35}12003 1200210140 11211032{135}1 4300202245 2332452012 2450014262
0102210020  ???? ???????  ???? ???????  ????220042 0020000000 412102?204 -0-
0302222 0400040120 2220100??0  ???? ???????  ???? ???????  ???? ??????? ???? ????????
??????????  ???? ???????  ???? ???????  ???? ???????  ???? ???????
??????????  ???? ???????  ???? ???????  ???? ???????  ???? ???????
??????????  ???? ???????  ???? ???????  ???? ???????  ?????
'Teilhardina asiatica'             1?0070010? 1?00?00003 0202042022 0232612002
1200030220 0020004560 4200202044 2422452002 2440023261 00022100?0
??????????  ?2?0?00001 2102220040 1000000000 2122020204 -0-0002222
0200041000 22200000?? 02????????  ???? ???????  ???? ???????  ???? ???????
??????????  ???? ???????  ???? ???????  ???? ???????  ???? ???????
??????????  ???? ???????  ???? ???????  ???? ???????  ???? ???????
??????????  ????2??2??  ???? ???????  ?1?1??1??  ???? ???????  ?????
'Teilhardina belgica'             11005??1??  ??00??????3 12???40002 0222512002
1200040220 0020005340 4300202044 2422452002 2440023261 0002210010
??????????  ???? ???????  ????220042 0000001000 2121020104 -0-0002222
0200040000 2220000???  ???? ???????  ???? ???????  ???? ??????? 2240020443
402625{35}144 2000010202 1200000000 1100000??? ?11?310??? ??1?01????
??????????  ???? ???????  ??10??????  ???? ???????  ???? ???????
??????????  ???? ???????  ???? ???????  ???? ???????  ?????
'Tetonius sp.'                   1102804441 1440?00010 2202040022 0221412003
1201120131 4421002502 4300202{13}44 2332452102 3440035261 0122210020
?344?322?0 ?210002?12 3102220041 0010100000 4124020104 -0-0202222
0400040220 2210200??2  ???? ???????  ???? ???????  ????220020243
402425{35}244 3021010202 1??1000100 101?110??? ???? ??????? ???? ???????
```
?????????? ?????????? ?????????? ?????????? 0 ?0??41??1 ?2???????? ??000???1
20????2?0? ?11??02??? ??111??1?? ??1???????? ????

'Varecia variegata' 1010414241 1200011034 2208041022 2224612022

1324320201 0000008268 0-020202-6 23020-2022 1-600180-2 2122210020
0120040020 4014000211 2102000040 0000000000 30240120-4 -0-0020002

0004040042 1300000000 0123420430 1022420230 1243112031 1420330044

4048431226 0000200030 0111001000 210122200? 0112321210 2210010224

0102440200 0311002001 2210004210 5100000002 4001020010 002000?120

101000?011 0102221020 0001100?1{04} 0032304010 000?001?0? ???1

'Wadilemur elegans' 1??????2?? ?????????? 2206030022 2202810--3

0200031402 2040028052 0-022022-6 24020-2212 2-600040-1 2002210010
?????????? ?????????? ?????????4? 000?0002?0 412?0?0200 2400000002

0002001004 0?????0??? ?????????? ?????????? ?????????? ?????????? ?????

?????????? 22 1241120?0? 02????????? ?????????? ?????????? ?????????? ?????

'Washakius insignis' 100040?0?1 1400?000?? 2213022222 0212612210

2303110020 2431203442 4110232423 2320422202 3230020222 2002410044
?????????? ????212 10?2242040 2020010200 2122220302 20-0224002

2401220000 0240400??? ????2????? ????2????? ????2????? ?340021343

4024246044 2020100212 1101000000 2010110??? ????2????? ????2????? ?????

????????? ????2????? ????2????? ????2????? ?????

'Xanthorhysis tabrumi' ??0??????? ?????????? 22???40022

0222212004 -202220220 4221208462 4100200024 2442422002 2240004421
2022210020 ????2????? ????2????? ????2????? ????2????? ?????

????????? ????2????? ????2????? ????2????? ????2????? ?????

????????? ????2????? ????2????? ????2????? ????2????? ?????

'Yunnanadapis folivorus' ??0??????? ???0?20203 2000040022 0226614222

1204020220 0000208458 0-00042200 24220-2222 20030240-1 4001210000
????????? ????2????? ????2????? 0 002022?000 20?2420200 00-000001

24?2221240 24004?0??? ????2????? ????2????? ????2????? ????2????? ?????

????????? ????2????? ????2????? ????2????? ????2????? ?????

END;

ctype ord : 1-4 5 7-9 12-15 18 21 22-28 30-35 37-61 63-67 69-75 77-78 80-84

86-88 91-95 98 99-100 102-104 106-109 111-115 117-118 120 123-131 133 135-136 138 140-141 143-162 164-166 168-180 182-187 189 191 193-197 199-200

202-204 206-207 209 212-221 223-231 233 235 238-240 242 244 246-248 251

254-255 257-258 264-266 268 271-272 279-280 282-286 288-290 292-293 296-298

300-302 304 307-309 311 313 316 318-322 324-326 328 333 338-339

347 353-355 357 359 363 367 369-373 376-377 380-382;

wts 1 : all;
wts 0.5 : 5 7-9 12-14 18 21-23 24-28 30-35 37-40 42-53 55-61 63-67 69-75 77

78 80-83 86-88 91-93 95 98-100 102-104 106-107 111-112 114 117-118 123-131

133 135-136 138 140-141 144-162 164-166 168-180 183-187 189 191 193-197

199-200 202 -204 209 212 -221 223 -231

240 248 264 279-280 285-286 288-289 292-293 296-298 300-302 307 309 311 316

318-322 324-326 333 338 347 354-355 357 359 367 369-373 376-377 380-381;
Character-taxon-matrix

Analysis 2: We use a matrix with 38 taxa and 326 morphological characters based on Jaeger
et al. (2020).

#NEXUS

[ File saved by NDE version 0.5.0, Thu Feb 15 17:53:16 2024 ]

BEGIN TAXA;

DIMENSIONS NTAX=38;

TAXLABELS

'Donrussellia gallica+sp.'

'Notharctus robustior'

'Adapis parisiensis'

'Cantius eppsi'

'Leptadapis magnus'

'Absarokius abotti+sp.'

'Hemiacodon gracilis'

'Shoshonius cooperi'

'Teilhardina americana'

'Xanthorhysis tabrumi'

'Tarsius spp.'

'Eosimias spp.'

'Phenacopithecus spp.'

'Bahinia pondaungensis'

'Proteopithecus sylviae'

'Serapia eocaena'

'Apidium phiomense'

'Parapithecus fraasi'

'Qatrania wingi'

'Simonsius grangeri'

'Biretia spp.'

'Dolichocebus gaimanensis'

'Branisella boliviana'

'Callicebus moloch'

'Saimiri sciureus'

'Saguinus spp.'

'Aotus spp.'

'Myanmarpithecus yarshensis'

'Ganlea megacanina'

'Pondaungia cotteri'

'Siamopithecus eocaenus'

'Aegyptopithecus zeuxis'

'Taqah propliopithecid'

'Catopithecus browni'

'Oligopithecus savagei'

'Oligopithecus rogeri'

'Propliopithecus spp.'

'Propliopithecus chirobates'

;

ENDBLOCK;

BEGIN CHARACTERS;

DIMENSIONS NCHAR=326;

FORMAT DATATYPE=STANDARD MISSING=? GAP=- SYMBOLS="012345";

CHARLABELS
[1] 'i1. Lower incisor number'
[2] 'i2. Lower incisor occlusal arrangement'
[3] 'i3. Lower incisor crown spacing'
[4] 'i4. I/2-C diastema'
[5] 'i5. I/1-2 size (ratio of I/1-2 area to M/1 area)'
[6] 'i6. I/1:1/2 proportions (ratio of I/1 area to I/2 area)'
[7] 'i7. I/1 crown width (spatulate incisors only)'
[8] 'i8. I2 crown cross-sectional shape (ratio of m-d length to b-1 breadth)'
[9] 'i9. Lower incisors crown height (crown heights judged from cementoenamel junction to crown tip on the buccal surface)'
[10] 'i11. Lower incisor roots'
[11] 'i12. Lower incisor crowns'
[12] 'i14. I/1 crown shape'
[13] 'i15. I/2 heel development (a lingual swelling at the base of crown)'
[14] 'i17. Lower first incisor lingual cingulum.'
[15] 'i19. Relative size of I1 to M1 (based on occlusal areas).'
[16] 'c1. Female C1 cross-sectional area relative to molar cross sectional area.'
[17] 'c2. C1/1 dimorphism (square root of male C1 area/square root of female C1 area).'
[18] 'c3. C/1 cross-sectional shape.'
[19] 'c4. C/1 lingual crest development.'
[20] 'c5. Canine paracristid (not scored if species has canine incorporated into a tooth comb).'
[21] 'c6. Canine height (females).'
[22] 'ML18. Lower canine crown.'
[23] 'ML19. Lower canine root.'
[24] 'p1. P1/1'
[25] 'p2. P/2.'
[26] 'OC1. Relative size (MDxBL) of p2 and p3'
[27] 'OC2. Relative size (MDxBL) of p2 and P2'
[28] 'p3. P/2 roots.'
[29] 'p4''. P/3 roots'
[30] 'p4". P/4 roots.'
[31] 'p5. Premolar crowding (overlapping of crowns).'
[32] 'p6. P/3 paraconid'
[33] 'OC7. P/3 paraconid height'
[34] 'p7. P/4 paraconid.'
[35] 'p9''. P/4 paraconid position (labiolingually)'
[36] 'p9'''. P/4 paraconid position (mesiodistally)'
[37] 'p11. P/3-4 cristid obliqua'
[38] 'p13. P/2 protoconid height and shape'
[39] 'p14. P/4 metaconid position'
[40] 'p15. P/2 metaconid size'
[41] 'p16. P/3 metaconid size'
[42] 'p17. P/4 metaconid size'
[43] 'p18. P/4 trigonid-configuration of lingual wall'
[44] 'p19. P/3 entoconid and lingual talonid crest'
[45] 'p20. P/4 entoconid and lingual talonid crest: '
[46] 'p21. P/4 lateral and medial protocristid'
[47] 'p22. P/3 lateral protocristid orientation'
[48] 'p23. P/4 lateral protocristid orientation'
[49] 'p24. P/3-4 posterior trigonid wall'
[50] 'p25. P/3-4 hypoconid size'
[51] 'p26. P/3-4 hypoconid (or distal terminus of oblique cristid) position'
[52] 'p27. P/4 hypocristid shearing development: '
[53] 'p28. P/2 buccal cingulum development'
[54] 'p29. Lower premolar inflation'
[55] 'p30. P/4 exodaenodonty'
[56] 'p31. P/4 talonid length (ratio of midline m-d length of trigonid to m-d length of talonid)'
[57] 'p33. Premolar orientation'
[58] 'OC 8. Lower premolar crown configuration'
[59] 'p34. P/4 anterobuccal cingulum development'
[60] 'p36. P/4 postprotoconid ridge'
[61] 'p37. P/4 postmetaconid ridge'
[62] 'p40. P/4 paraconid height'
[63] 'p41. P3-4 protoconid height'
[64] 'p42. P/3 to P/4 area'
[65] 'p43. P/4 m-d L/ b-1 W'
[66] 'p44. Ratio of P/4 area to M/1 area'
[67] 'p45a. P/3 root orientation'
[68] 'p45b. P/4 root orientation'
[69] 'm4. M/3 root number'
[70] 'm6. M/2 trigonid width (ratio of buccolingual breadths of trigonid and talonid)'
[71] 'm7. M/3 trigonid width (based on relative buccolingual breadths)'
[72] 'OC3. M/1 paraconid'
[73] 'm8''. M/1 paraconid position '
[74] 'm9''. M/2 paraconid position '
[75] 'OC4. M/3 paraconid'
[76] 'm10. M/3 paraconid position '
[77] 'm8910. M/2-3 paraconid location'
[78] 'm11. M/1 parastylid (= premetacristid)'
[79] 'm12. Molar metastylids (postmetacristids)'
[80] 'm13. M/3 hypoconulid'
[81] 'm14. M/3 heel'
[82] 'm15. Molar enamel surface'
[83] 'm16. M/1 trigonid height (ratio of trigonid height to talonid height measured on the buccal aspect of the crown)'
[84] 'm17. M/1-2 cusp relief'
[85] 'm18. M/1 trigonid lingual configuration'
[86] 'm19. M/1 metaconid position'
[87] 'm20. M/2 paraconid development'
[88] 'm21. M/1-2 lateral protocristid orientation'
[89] 'm22. M/1 distal trigonid wall'
[90] 'm23. M/2 distal trigonid wall'
[91] 'm24. M/1-3 wear facet X'
[92] 'm25. M/1-2 entoconid'
[93] 'm26. M/1-2 postentoconid sulcus'
[94] 'm27. M/1 hypoconulid size'
[95] 'm28. M/2 hypoconulid size'
[96] 'm29. M/3 hypoconulid size'
[97] 'm30. M/1-2 hypoconulid position'
[98] 'm31. M/1-2 cristid obliqua development'
[99] 'm32. M/1 cristid obliqua orientation'
[100] 'm33. M/2 cristid obliqua orientation'
[101] 'm34. M/1 cristid obliqua terminus'
[102] 'm35. M/2 cristid obliqua terminus'
[103] 'm36. M/3 cristid obliqua terminus'
[104] 'm37. M/1-2 centroconid development'
[105] 'm38. M/1-2 hypocristid development'
[106] 'm39. M/3 hypocristid development'
[107] 'm40. Lingual configuration of M/1-2 talonid'
[108] 'm41. M/1-2 distal fovea'
[109] 'm42. M/1-2 hypocristid configuration'
[110] 'm43. M/1-2 cristid obliqua'
[111] 'm44. Molar cusp inflation'
[112] 'm45. M/1-2 buccal cingulum development'
[113] 'm46. M/1 hypoflexid depth'
[114] 'm47. M/2 hypoflexid depth'
[115] 'm53. Ratio of M/2 length to M/3 length'
[116] 'm55. M/1 mesiodistal length/buccolingual breadth'
[117] 'm56. Convergence of buccal and lingual molar cusp walls'
[118] 'm57. M/1-2 entoconid position relative to hypoconid'
[119] 'ML88. M/1-3 Pre-entocristid'
[120] 'I1. I1/-I2/ interstitial contact'
[121] 'I2. I1/-I1/ interstitial contact'
[122] 'I3. I2/-C diastema'
[123] 'I4. I1/ area:I2/ area'
[124] 'I5. I1/ size (I1/ area: M1/ area)'
[125] 'I6. I1/ occlusal shape (mesiodistal length/buccolingual breadth)'
[126] 'I7. I2/ occlusal shape (mesiodistal length/buccolingual breadth)'
[127] 'I10. I1/ occlusal edge orientation (for spatulate incisors only; all others scored as " ? ")'
[128] 'I11. I1-2/ lingual cingulum'
[129] 'I12. I1/ basal lingual cusp'
[130] 'I13. I1/-I2/ buccal cingulum'
[131] 'C1. C1/ cross-sectional shape'
[132] 'C2. Upper canine occlusion'
[133] 'C3. C1/ mesial groove (females)'
[134] 'C4. C1/ lingual cingulum'
[135] 'OC5. P2/"
[136] 'P1". P2/ root number'
[137] 'P2. P3/ root number'
[138] 'P3. P4/ root number'
[139] 'P4". Ratio of P2/ area to P3/ area'
[140] 'P5. Ratio of P4/ area to M1/ area'
[141] 'P6. P2/ occlusal outline'
[142] 'OC6. P3/ occlusal outline'
[143] 'P7". P4/ occlusal outline'
[144] 'P8. P3-4/ trigon/talon proportions'
[145] 'P9. P3/ protocone'
[146] 'P10. P4/ metacone'
[147] 'P11. P4/ protocone'
[148] 'P12". P2/ protocone'
[149] 'P13". P2/ hypocone'
[150] 'P14. P4/ paraconule'
[151] 'P15. P3-4/ parastyles'
[152] 'P16. P3-4/ metastyles'
[153] 'P17. P3-4/ postprotocrista'
[154] 'P18". P2-4/ distal crown margin'
[155] 'P19. P3-4/ lingual cingulum'
[156] 'P20. P3/ metacone'
[157] 'P21. P3-4/ buccal cingulum development'
[158] 'ML126. P4/ hypocone'
[159] 'ML127. P3/ hypocone'
[160] 'M1. M1-2/ root number'
[161] 'M2. M3/ root number'
[162] 'M3. M2/ shape (bl/md)'
[163] 'M4. Ratio of M1/ area to M2/ area'
[164] 'M7". M1-2/ metaconule'
[165] 'M9. M1-2/ preprotoconule'
[166] 'M10. M1/ hypocone size'
<table>
  <tbody>
    <tr>
      <td>[167]</td>
      <td>'M11. M2/ hypocone size'</td>
    </tr>
    <tr>
      <td>[168]</td>
      <td>'M12''. M1-2/ hypocone position'</td>
    </tr>
    <tr>
      <td>[169]</td>
      <td>'M13. M1-2/ prehypocrista development'</td>
    </tr>
    <tr>
      <td>[170]</td>
      <td>'M14. M3/ prehypocrista development'</td>
    </tr>
    <tr>
      <td>[171]</td>
      <td>'M15. M1/ or M2/ paraconule position'</td>
    </tr>
    <tr>
      <td>[172]</td>
      <td>'M16. M1-2/ metaconule'</td>
    </tr>
    <tr>
      <td>[173]</td>
      <td>'M17'''. M1-2/ mesostyle size'</td>
    </tr>
    <tr>
      <td>[174]</td>
      <td>'M17". M1-2/ mesostyle position'</td>
    </tr>
    <tr>
      <td>[175]</td>
      <td>'M20. P4/-M1/ pericone'</td>
    </tr>
    <tr>
      <td>[176]</td>
      <td>'M22. M1-3/ lingual cingulum development'</td>
    </tr>
    <tr>
      <td>[177]</td>
      <td>'M24. M1-2/ buccal cingulum development'</td>
    </tr>
    <tr>
      <td>[178]</td>
      <td>'M27. M1-2/ pre-metaconule cristae'</td>
    </tr>
    <tr>
      <td>[179]</td>
      <td>'M28. M1-2/ post-metaconule cristae'</td>
    </tr>
    <tr>
      <td>[180]</td>
      <td>'M30. M3/ paraconule'</td>
    </tr>
    <tr>
      <td>[181]</td>
      <td>'M31. Molar protocone lingual inflation'</td>
    </tr>
    <tr>
      <td>[182]</td>
      <td>'M33. M2/ buccal expansion of paracone (specify which tooth)'</td>
    </tr>
    <tr>
      <td>[183]</td>
      <td>'M34. M3/ metacone'</td>
    </tr>
    <tr>
      <td>[184]</td>
      <td>'M36. M3/ hypocone'</td>
    </tr>
    <tr>
      <td>[185]</td>
      <td>'M37. M1/ paraconule size'</td>
    </tr>
    <tr>
      <td>[186]</td>
      <td>'M44. M1-3/ anterior cingulum'</td>
    </tr>
    <tr>
      <td>[187]</td>
      <td>'M46. M3/ size relative to M1/'</td>
    </tr>
    <tr>
      <td>[188]</td>
      <td>'ML147. M1-2/ metastyle'</td>
    </tr>
    <tr>
      <td>[189]</td>
      <td>'ML148. M1-2/ parastyle'</td>
    </tr>
    <tr>
      <td>[190]</td>
      <td>'ML149. M1-2/ parastyle position'</td>
    </tr>
    <tr>
      <td>[191]</td>
      <td>'ML150. M1-2/ metastyle position'</td>
    </tr>
    <tr>
      <td>[192]</td>
      <td>'ML151. M1-3/ posterior cingulum'</td>
    </tr>
    <tr>
      <td>[193]</td>
      <td>'ML152. M1-3/ posterior margin (waisted between buccal and lingual cusps)'</td>
    </tr>
    <tr>
      <td>[194]</td>
      <td>'ML153. M1-2/ postparacrista'</td>
    </tr>
    <tr>
      <td>[195]</td>
      <td>'ML154. M1-2/ premetacrista'</td>
    </tr>
    <tr>
      <td>[196]</td>
      <td>'ML155. M1-3/ protocone arrangement'</td>
    </tr>
    <tr>
      <td>[197]</td>
      <td>'ML156. M1-2/ postprotocrista development'</td>
    </tr>
    <tr>
      <td>[198]</td>
      <td>'ML157. M1/ postprotocrista length'</td>
    </tr>
    <tr>
      <td>[199]</td>
      <td>'ML158. M2/ postprotocrista length'</td>
    </tr>
    <tr>
      <td>[200]</td>
      <td>'ML159. M1/ postprotocrista direction'</td>
    </tr>
    <tr>
      <td>[201]</td>
      <td>'ML160. M2/ postprotocrista direction'</td>
    </tr>
    <tr>
      <td>[202]</td>
      <td>'ML161. M1/ postprotocrista terminus'</td>
    </tr>
    <tr>
      <td>[203]</td>
      <td>'ML162. M2/ postprotocrista terminus'</td>
    </tr>
    <tr>
      <td>[204]</td>
      <td>'ML163. M1-2/ preprotocrista'</td>
    </tr>
    <tr>
      <td>[205]</td>
      <td>'ML164. M1/ preprotocrista connection (buccal side)'</td>
    </tr>
    <tr>
      <td>[206]</td>
      <td>'ML165. M2/ preprotocrista connection (buccal side)'</td>
    </tr>
    <tr>
      <td>[207]</td>
      <td>'ML166. M1-2/ postparaconule crista'</td>
    </tr>
    <tr>
      <td>[208]</td>
      <td>'ML168. M1-2/ hypometacrista'</td>
    </tr>
    <tr>
      <td>[209]</td>
      <td>'ML169. M1-2/ hypoparacrista'</td>
    </tr>
    <tr>
      <td>[210]</td>
      <td>'MLN. Hypometaconulecrista'</td>
    </tr>
    <tr>
      <td>[211]</td>
      <td>'CR1. Transverse septum arising from the cochlear housing'</td>
    </tr>
    <tr>
      <td>[212]</td>
      <td>'CR2. Extent of pneumatization of anterior accessory cavity'</td>
    </tr>
    <tr>
      <td>[213]</td>
      <td>'CR3. Pneumatization of mastoid (from epitympanic recess?)'</td>
    </tr>
    <tr>
      <td>[214]</td>
      <td>'CR4. Presence or absence of perbullar pathway'</td>
    </tr>
    <tr>
      <td>[215]</td>
      <td>'CR5. Anteroposterior location of posterior carotid foramen in bulla'</td>
    </tr>
    <tr>
      <td>[216]</td>
      <td>'CR6. Mediolateral position of posterior carotid foramen in bulla'</td>
    </tr>
    <tr>
      <td>[217]</td>
      <td>'CR7. Ventrodorsal position of the carotid foramen in the bulla'</td>
    </tr>
    <tr>
      <td>[218]</td>
      <td>'CR8. Position of posterior carotid foramen relative to fenestra cochleae'</td>
    </tr>
  </tbody>
</table>
[219] 'CR9. Position of the internal carotid canal relative to the fenestra cochleae'

[220] 'CR10. Position of the portion of the internal carotid/promontory artery (or its accompanying nerves) lying on the promontorium anterior to the fenestra cochleae'

[221] 'CR11. Size of stapedial and promontory canals'

[222] 'CR12. Morphology of promontory canal, when present'

[223] 'CR14. Position of ventral edge of the tympanic bone'

[224] 'CR15. The shape of the tympanic bone'

[225] 'CR16. Morphology of annular bridge'

[226] 'CR17. Encroachment of the auditory bulla on the pterygoid fossa'

[227] 'CR18. Nature of contact between the lateral pterygoid plate and the bulla wall'

[228] 'CR19. Extent of contact between the lateral pterygoid plate and the bulla wall'

[229] 'CR20. Flange of basioccipital overlapping medial bulla wall'

[230] 'CR21. Suprameatal foramen'

[231] 'CR22. Patent parotic fissure'

[232] 'CR23. Size of orbits'

[233] 'CR24. Postorbital closure'

[234] 'CR25. Composition of the postorbital septum'

[235] 'CR26. Zygomatic-lacrimal contact'

[236] 'CR27. Pronounced interorbital constriction'

[237] 'CR28. Contact between lacrimal and palatine'

[238] 'CR29. Foramen rotundum'

[239] 'CR30. Position of lacrimal foramen'

[240] 'CR31. Metopic suture in adult'

[241] 'CR32. Orbital convergence'

[242] 'CR33. Posterior nasal spine'

[243] 'CR34. Posterior palatine torus'

[244] 'CR35. Pyramidal processes'

[245] 'CR36. Length of medial pterygoid plate'

[246] 'CR37. Snout length'

[247] 'CR38. Maxillary depth'

[248] 'CR39. Complete symphyseal fusion'

[249] 'CR40. Temporomandibular joint morphology'

[250] 'CR41. Entoglenoid process morphology'

[251] 'CR42. Inter-incisor diastema width'

[252] 'CR43. Coronoid height relative to condyle'

[253] 'CR44. Condyle height relative to toothrow'

[254] 'CR45. Corpus robusticity'

[255] 'CR46. Zygomatico-parietal contact at pterion'

[256] 'CR48. Epitympanic crest'

[257] 'CR49. Broad ascending wing of premaxilla'

[258] 'CR50. Basioccipital stem'

[259] 'CR51. Choanal shape'

[260] 'CR52. Orientation of the mandibular symphysis'

[261] 'H1. Shape of distal edge of the humeral trochlea'

[262] 'H2. Relative heights of medial and lateral edges of humeral trochlea'

[263] 'H3. Trochleocapitular ridge'

[264] 'H4. Waisted trochlea (Minimum trochlear diameter/maximum trochlear diameter x 100)'

[265] 'H5. Width of capitulum relative to trochlea (100 x ventral capitulum width/ventral trochlear width)'

[266] 'H6. Entepicondylar foramen'

[267] 'H7. Entepicondylar foramen position'

[268] 'H8. Medial epicondyle size'

[269] 'H9. Dorsal placement of medial epicondyle'
<table>
  <tbody>
    <tr>
      <td rowspan="10">trochlea</td>
      <td>[270] 'H10''</td>
      <td>Shape of the lateral edge of the dorsal</td>
    </tr>
    <tr>
      <td>[271] 'H10''</td>
      <td>Shape of the medial edge of the dorsal</td>
    </tr>
    <tr>
      <td>[272] 'H11. Dorsoepitrochlear fossa'</td>
      <td></td>
    </tr>
    <tr>
      <td>[273] 'H12. Olecranon fossa shape'</td>
      <td></td>
    </tr>
    <tr>
      <td>[274] 'H14. Brachialis flange'</td>
      <td></td>
    </tr>
    <tr>
      <td>[275] 'H15. Bicipital groove morphology'</td>
      <td></td>
    </tr>
    <tr>
      <td>[276] 'H18. Capitular tail'</td>
      <td></td>
    </tr>
    <tr>
      <td>[277] 'H19. Ratio of humerus length to femur length (H/F)'</td>
      <td></td>
    </tr>
    <tr>
      <td>[278] 'W2. Ulnar-pisiform articulation.'</td>
      <td></td>
    </tr>
    <tr>
      <td>[279] 'OP1. Gluteal tuberosity'</td>
      <td></td>
    </tr>
    <tr>
      <td>[280] 'OP2. Position of posterior gluteal tuberosity'</td>
      <td></td>
    </tr>
    <tr>
      <td rowspan="10">femur/ mediolateral diameter of distal femur).</td>
      <td>[281] 'F1. Length of femoral neck'</td>
      <td></td>
    </tr>
    <tr>
      <td>[282] 'F2. Angle of femoral neck'</td>
      <td></td>
    </tr>
    <tr>
      <td>[283] 'F3. Angle of lesser trochanter'</td>
      <td></td>
    </tr>
    <tr>
      <td>[284] 'F4. Size of third trochanter'</td>
      <td></td>
    </tr>
    <tr>
      <td>[285] 'F5. Knee index (Antero-posterior diameter of distal femur)'</td>
      <td></td>
    </tr>
    <tr>
      <td>[286] 'F6. Femoral head shape'</td>
      <td></td>
    </tr>
    <tr>
      <td>[287] 'F7. Anterior extension of greater trochanter'</td>
      <td></td>
    </tr>
    <tr>
      <td>[288] 'F8. Anterior bend of proximal femur'</td>
      <td></td>
    </tr>
    <tr>
      <td>[289] 'F9. Relative length of trochanteric fossa'</td>
      <td></td>
    </tr>
    <tr>
      <td>[290] 'F10. Presence of intertrochanteric crest'</td>
      <td></td>
    </tr>
    <tr>
      <td>[291] 'F11. Size of lesser trochanter'</td>
      <td></td>
    </tr>
    <tr>
      <td>[292] 'F12. Lateral rim of knee'</td>
      <td></td>
    </tr>
    <tr>
      <td>[293] 'T1''</td>
      <td>Fusion of tibia and fibula</td>
    </tr>
    <tr>
      <td>[294] 'T1''</td>
      <td>Articulation tibia/fibula</td>
    </tr>
    <tr>
      <td>[295] 'T3. Shape of distal surface of tibia'</td>
      <td></td>
    </tr>
    <tr>
      <td>[296] 'T4. Rotation of the medial malleolus'</td>
      <td></td>
    </tr>
    <tr>
      <td>[297] 'T5. Shape of medial malleolar articular surface'</td>
      <td></td>
    </tr>
    <tr>
      <td>[298] 'T6. Shape of distal tibial shaft'</td>
      <td></td>
    </tr>
    <tr>
      <td>[299] 'T7. Position of tibialis posterior groove'</td>
      <td></td>
    </tr>
    <tr>
      <td>[300] 'A1. Position of the flexor hallucis longus groove'</td>
      <td></td>
    </tr>
    <tr>
      <td>[301] 'A2''</td>
      <td>Shape of talo-fibular facet.</td>
    </tr>
    <tr>
      <td>[302] 'A4''</td>
      <td>Development of the talar posterior trochlear</td>
    </tr>
    <tr>
      <td rowspan="16">shelf'</td>
      <td>[303] 'A5''</td>
      <td>Talar neck length (NL/TL x 100)</td>
    </tr>
    <tr>
      <td>[304] 'A6. Medial talo-tibial facet'</td>
      <td></td>
    </tr>
    <tr>
      <td>[305] 'A7. Lateral talar trochlear asymmetry'</td>
      <td></td>
    </tr>
    <tr>
      <td>[306] 'A8. Talar cotylar fossa'</td>
      <td></td>
    </tr>
    <tr>
      <td>[307] 'A9. Width of the head of the talus (HW/HHT x 100)'</td>
      <td></td>
    </tr>
    <tr>
      <td>[308] 'GEB1. Talar neck angle'</td>
      <td></td>
    </tr>
    <tr>
      <td>[309] 'GEB2. Talar body height (HT/MTRW x 100)'</td>
      <td></td>
    </tr>
    <tr>
      <td>[310] 'GEB3. TW/TL x 100'</td>
      <td></td>
    </tr>
    <tr>
      <td>[311] 'CA1. Anterior calcaneal elongation'</td>
      <td></td>
    </tr>
    <tr>
      <td>[312] 'CA2. Position of the peroneal tubercle'</td>
      <td></td>
    </tr>
    <tr>
      <td>[313] 'CA3. Posterior calcaneal bowing'</td>
      <td></td>
    </tr>
    <tr>
      <td>[314] 'CA4. Calcaneo-cuboid articulation'</td>
      <td></td>
    </tr>
    <tr>
      <td>[315] 'N1. Length relative to width'</td>
      <td></td>
    </tr>
    <tr>
      <td>[316] 'N3. Morphology of the naviculocuboid articulation'</td>
      <td></td>
    </tr>
    <tr>
      <td>[317] 'E1. Shape of Entocuneiform/MT1 articulation'</td>
      <td></td>
    </tr>
    <tr>
      <td>[318] 'E2. Lateral process of entocuneiform'</td>
      <td></td>
    </tr>
    <tr>
      <td>[319] 'O1. Foot axis'</td>
      <td></td>
    </tr>
    <tr>
      <td>[320] 'O2. Toilet claw'</td>
      <td></td>
    </tr>
    <tr>
      <td>[321] 'O3. Prehallux'</td>
      <td></td>
    </tr>
    <tr>
      <td>[322] 'O4. Metatarsus length'</td>
      <td></td>
    </tr>
    <tr>
      <td>[323] 'MT1. Peroneal tubercle of MTI'</td>
      <td></td>
    </tr>
    <tr>
      <td>[324] 'MT2. Hallux length'</td>
      <td></td>
    </tr>
    <tr>
      <td>[325] 'V1. Optic fovea'</td>
      <td></td>
    </tr>
    <tr>
      <td>[326] 'V2. Tapetum lucidum'</td>
      <td></td>
    </tr>
  </tbody>
</table>
; STATELABELS

1

'three '
'two '
'one I/1 present, I/2 absent '
'lower incisors absent ',

2

arcade) '

'Arcuate battery from lateral perspective (U-shape
'cusp tips staggered (V-shaped arcade) ',

3

'No spaces '
'Spaces present between crowns ',

4

'present '
'absent ',

5

'very small (<0.69) '
'moderate sized (<0.70, <1.07) '
'large (>1.07) ',

6

'I/1 much smaller than I/2 (<0.65) '
'I/1 smaller than I/2 (>0.65, <0.82) '
'I/1 almost as large as I/2 (>0.83, <1.00) '
'I/1>I/2 (>1.01, <1.25) '
'I/1>>I/2 (>1.25) ',

7

'considerably wider (m-d) than root (spatulate) '
'narrow at apex, wider than root '
'"styliform" (crown apex approximately the same

width as the cervical margin)',

8

'rounded oval (> 0.64) '
'mesiodistally compressed (< 0.64) ',

9

'low crowned '
'moderately high crowned '
'high crowned ',

10

'erect or vertical '
'slightly procumbent '
'very procumbent ',

11

'erect or vertical'
procumbent
'very procumbent ',

12

spatulate
'lanceolate, pointed ',

13

'heel absent '
'heel present ',

14

'absent to weak '
'strong but incomplete '
'strong and complete ',

15

'I/1 very small (I/1 << M/1) '
'moderately enlarged (I/1 < or = M/1) '
'grossly enlarged (I/1 > M/1) ',
16
' very small (C1/M1 < 0.40) '
' moderate (> 0.4, < 0.80) '
' large (> 0.80, < 1.20); very large (> 1.20) ',
17
' low (< 1.07) '
' moderate (> 1.07, < 1.17) '
' high (>1.17) ',
18
' rounded oval '
' mesiodistally compressed '
' buccolingually compressed ',
19
' rounded '
' sharp ',
20
' oblique to occlusal plane '
' nearly horizontal to occlusal plane '
' forms part of cropping mechanism with I/1-2 ',
21
' low, squat '
' narrow, short '
' tall, at or above tooth row ',
22
' erect or vertical '
' procumbent '
' very procumbent ',
23
' erect or vertical '
' slightly procumbent '
' very procumbent ',
24
' present '
' absent ',
25
' present '
' absent ',
26
' p2<<p3 (ratio<0.6) '
' p2<p3 (0.6<ratio<0.9) '
' p2> or =p3 (ratio>0.9) ',
27
' P2 larger or equal to p2 '
' P2 smaller than p2 ',
28
' single '
' double ',
29
' single '
' double ',
30
' single '
' double ',
31
' no crowding '
' slightly crowded '
' very crowded-mesial root positioned buccal to
    distal root ',
32
' large '
' small '
'absent or extremely small ',
33
Low
moderate
'High (nearly as high as the protoconid)',
34
'large '
'small '
'absent or extremely small ',
35
'mesial to protoconid'
'mesiolingual, between protoconid and metaconid'
'mesial to metaconid',
36
'widely spaced from the metaconid '
'twinned with metaconid ',
37
'absent '
'weak '
'strong ',
38
'projects above protoconids of P/3-4 '
'massive, projects above protoconids of P/3-4 '
'not projecting, in line with P/3 '
'extremely short, shorter than P/3 ',
39
'close to protoconid '
'widely spaced from protoconid ',
40
'absent or trace '
'small ',
41
'absent or trace '
'small '
'large (as big as protoconid) ',
42
'absent or trace'
'small '
'large (as big as protoconid) ',
43
'closed '
'open ',
44
'absent '
'lingual talonid crest present but an entoconid does
not stand out above it '
'entoconid forms a small discrete cusp ',
45
'absent '
'lingual talonid crest present but an entoconid does
not stand out above it '
'entoconid forms a small discrete cusp ',
46
'continuous between metaconid and protoconid '
'discontinuous between metaconid and protoconid ',
47
'transversely oriented '
'distolingually oriented '
'absent ',
48
'transversely oriented '