A new pliopithecoid genus from the early Miocene of Uganda

Journal of Human Evolution 50 (2006) 568e586

A new pliopithecoid genus from the early Miocene of Uganda

James B. Rossie a,*, Laura MacLatchy b

a Section of Vertebrate Paleontology, Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, PA 15213, USAb Department of Anthropology, University of Michigan, 550 East University Avenue, Ann Arbor, MI 48109, USA

Received 17 June 2005; accepted 23 December 2005

Abstract

A partial face and mandible from the early Miocene site of Napak IX in Uganda are described here as a new genus and species of catarrhineprimate, Lomorupithecus harrisoni gen. et sp. nov. The face is among the most complete specimens known for a Miocene small-bodied catar-rhine. Several aspects of its anatomy indicate that the new species is a stem catarrhine, and as such, it may provide valuable information pertain-ing to the primitive catarrhine cranial morphotype. Lomorupithecus is most similar in its facial anatomy to members of the Pliopithecoidea, andthese similarities could be interpreted in three ways. They could be symplesiomorphies, which would support the traditional view of the prim-itive catarrhine cranial morphotype; they could be synapomorphies reflecting a phylogenetic position of Lomorupithecus within Pliopithecoidea;or they could represent convergence. Phylogenetic analysis of Lomorupithecus along with 35 other primates indicates that it is a pliopithecoid.As such, it would be the oldest and only Afro-Arabian member of this otherwise Eurasian clade.� 2006 Elsevier Ltd. All rights reserved.

Keywords: Pliopithecoid; Uganda; Miocene; Napak; Catarrhine

Introduction

In 2002, joint paleontological work by the Uganda NationalMuseum, Makerere University, and Boston University at theearly Miocene site of Napak IX in Uganda led to the discoveryof a partial face and juvenile hemi-mandible of a small catar-rhine primate. The facial anatomy of small-bodied Miocenecatarrhines is virtually unknown (Harrison, 1982, 2002), andthis lack of evidence has surely contributed to the ambiguityof their phylogenetic interrelationships. Here we describethis material as a new genus and species and conduct a phylo-genetic analysis to investigate its relationship to othercatarrhines.

* Corresponding author. Department of Anthropology, Stony Brook Univer-

sity, SBS Building, Stony Brook, NY 11794, USA. Tel.: þ1 631 632 1843.

E-mail addresses: [email protected] (J.B. Rossie), maclatch@

umich.edu (L. MacLatchy).

0047-2484/$ - see front matter � 2006 Elsevier Ltd. All rights reserved.

doi:10.1016/j.jhevol.2005.12.007

Systematics

Order Primates Linnaeus, 1758Suborder Anthropoidea Mivart, 1864Infraorder Catarrhini Geoffroy, 1812Superfamily Pliopithecoidea Zapfe, 1960Family incertae sedisLomorupithecus gen. nov.

Generic diagnosis. Differs from all other Miocene catar-rhines (including pliopithecoids) in the combination of ex-treme mesiodistal brevity of the upper premolar crowns, theovoid and symmetrical shape of the P4 crown, the twinningof the M2 hypoconid and hypoconulid, the expansion of theM2 distal fovea, and the prowlike paracristid. It differs fromall other African Miocene catarrhines for which the regionis preserved (Proconsul, Turkanapithecus, Afropithecus; Rae,1999) in retaining a platyrrhine-like configuration of the con-tact between the premaxillae and nasal bones. Lomorupithecusdiffers from Dendropithecus, Limnopithecus, Kalepithecus,

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569J.B. Rossie, L. MacLatchy / Journal of Human Evolution 50 (2006) 568e586

and Simiolus in having the protoconid mesial to the metaconidon M1 and M2; from Dendropithecus and Micropithecus inhaving less heteromorphic P3 cusps; from Kalepithecus andDendropithecus in having a more sloping lingual face of P3;from Dendropithecus and Simiolus in having a less pro-nounced P4 lingual cingulum, and less buccolingually com-pressed upper canines; from Simiolus in having no P3

lingual cingulum; from Limnopithecus and Kalepithecus inlacking an evenly convex distal margin of the P3 crown;from Kalepithecus, Micropithecus, and Simiolus in havinga mesiodistally shorter M1; from Kogolepithecus (Pickfordet al., 2003) in lacking a bifid metaconid and entoconid, andretaining a hypometacristid on M2; from Laccopithecus, Epi-pliopithecus, and Pliopithecus antiquus in having a lessobliquely oriented cristid obliqua; and from Micropithecusin having a pronounced mesiolingual cingulum on M1, a molartrigon considerably wider than long, rounded, barlike (as op-posed to sharp) inferior orbital rim, and orbits positionedhigher on the face. Lomorupithecus differs from all nyanzapi-thecines in having buccolingually broader upper premolarsand M1, and from all cercopithecoids in lacking bilophodontmolars.

Type species. L. harrisoni.Etymology. Combination of the Greek pithekos, meaning

‘‘ape’’ or ‘‘trickster,’’ and the Karimojong word lomoru, mean-ing ‘‘mountain man,’’ in reference to Mount Akiism, on whichthe site is located, as well as in honor of the late Will Downs,who was named ‘‘Lomoru’’ by the local Karimojong.

L. harrisoni sp. nov.

Holotype. BUMP1 266, partial face preserving maxilla, pre-maxilla, nasal bones, crowns of left P3eM1, and partial crownsof left canine and right P3eM1.

Hypodigm. Type specimen and BUMP 268, a juvenile man-dible containing M1, M2 (exposed, but unerupted), roots ofdm2, and alveoli for I1, I2, dc, and dm1.

Specific diagnosis. As for genus; a species with a body massof approximately 4.3 kg based on M1 size (Conroy, 1987).

Etymology. Named for Professor Terry Harrison in recogni-tion of his contributions to the study of Miocene catarrhineevolution.

Horizon and locality. Locality IX of the Napak sequence,early Miocene of Uganda. Napak localities I, IV, V, and IXare intercalated with volcanic tuffs (Bishop, 1964; Bishopet al., 1969; Pickford, 1981), and biotites from these tuffs atNapak I have produced dates of 19.5� 2 Ma and18.3� 0.4 Ma (adjusted for new constants; Ness et al.,1980). Petrological evidence suggests that the younger dateis discrepantly low (Bishop, 1964; Bishop et al., 1969), andthe fossiliferous sites are now believed to be between 19 and20 Ma (Senut et al., 2000).

1 Abbreviations are as follows: BUMP, Boston University/Uganda Museum/

Makerere University Paleontology Expeditions; UMP, Uganda Museum

of Paleontology; KNM, Kenya National Museums.

Comparative description

BUMP 266

Specimen BUMP 266 is a partial face consisting of thepalate anterior to the M1/M2 junction, as well as most of thepremaxilla, maxilla, lacrimal bones, and nasal bones (Fig. 1).The face was found in three pieces, all within a 2-m2 areaof outcrop. The palate was split in two near the midline, andthe interorbital fragment including the nasals and part of thepremaxilla was found as a separate piece down slope fromthe other two. The frontal process of the maxilla and the supe-rior tip of the ascending wing of the premaxilla cross the breakbetween the interorbital fragment and lower face on the rightside. There is a good fit between the two pieces, and the taper-ing of the premaxilla as it crosses the break demonstrates thatno interposing bone was lost. The specimen is an adult, judg-ing from the amount of wear on the M1s, and a male, based onthe lack of waisting of the canine root at the cemento-enameljunction, which is usually present in female catarrhines(Kelley, 1986). Because the facial morphology of small-bodiednon-cercopithecoid Miocene catarrhines is known from onlyMicropithecus clarki (Fleagle, 1975), Epipliopithecus vindobo-nensis (Zapfe, 1960), Pliopithecus zhanxiangi (Harrison et al.,1991), and Laccopithecus robustus (Pan, 1988), the followingtext will compare Lomorupithecus with fossil and extant taxatogether. Some comparisons to the stem cercopithecoidVictoriapithecus will also be included because of its relevanceto the ancestral catarrhine morphotype (see Benefit andMcCrossin, 1993).

Orbital region. A 5-mm-long portion of the inferior orbitalrim is preserved on the left side, just medial to the infraorbitalforamen and lateral to the lacrimal fossa, part of which is pre-served on the internal surface of the maxilla (Fig. 1a). This rimis a rounded bar that is broken laterally just at the point atwhich it would be met by its complementary process to en-close the infraorbital foramen. The arc of the rim impliesthat a considerable portion of the orbit lay laterally to it.The orbit was low, with the rim above the infraorbital foramenpositioned 17.7 mm above the P4eM1 alveoli. The most ante-rior portion of the rim is placed far forward, above the P3/P4

boundary.The lacrimal bone and its fossa are preserved on the right

side (Fig. 1b), as is the inferomedial portion of the infraorbitalcanal (Fig. 1d). The short broken surface between the two isthe same size as the contralateral rim fragment, which permitsa good reconstruction of the relationships among these ele-ments. The orbital rim (preserved on the left side) wouldmeet a steeply inclined lacrimal (preserved on the rightside), which formed a sharper inferomedial border to the orbit.The arc formed by these elements implies a wide interorbitalregion, which is confirmed by the nasal bones (describedbelow). The infraorbital canal is undivided and very short,as in gibbons, consisting of little more than a large foramenenclosed by a narrow, barlike process of bone. In gibbons,juvenile and subadult specimens reveal that the infraorbitalcanal/foramen is produced by the development of a barlike

570 J.B. Rossie, L. MacLatchy / Journal of Human Evolution 50 (2006) 568e586

Fig. 1. Craniodental remains of Lomorupithecus: anterior (a), posterior (b), palatal (c), and lateral (d) views of BUMP 266. Sutures between the nasal, premaxilla,

and maxilla in (a) have been accentuated for clarity. Abbreviations are as follows: fs, frontal sinus; ioc, infraorbital foramen; i1, central incisor; i2, lateral incisor;

lac, lacrimal crest; mat, matrix; ms, maxillary sinus; nld, nasolacrimal duct; orb, orbital rim; pmx, portion of ascending wing of premaxilla that overlaps nasal bone.

process of the inferior orbital margin that grows medially tomeet the frontal process of the maxilla as the bone remodelsaround the inferiorly drifting infraorbital canal (Fig. 2a). Asin gibbons, some colobines, Micropithecus clarki, Epipliopi-thecus, and Laccopithecus, the foramen lies medial and poste-rior to the lowest part of the orbital rim. The infraorbital canalof other anthropoids (e.g., great apes, cercopithecines, andVictoriapithecus) lies in a more lateral position relative tothe lacrimal canal, and traverses a more substantial portionof the maxilla, often including the maxillary sinus (Fig. 2c,d).

Interorbital region. Both nasal bones are nearly completeand remain attached to the most superior portions of the max-illa and premaxilla on the right side, and the maxilla alone onthe left side. The nasals are short and wide like those of livinggibbons, but in contrast to living hominoids, their width doesnot expand inferiorly around the lateral aspect of the nasal ap-ertures. Together, the two nasals form a diamond shape (withtruncated superior and inferior ends), similar to the nasals ofsmall orthognathic platyrrhines with wide interorbital regions,such as Callicebus. Unlike Callicebus, however, the lacrimal


fossa is well inferior to the nasofrontal suture. The nasals ofTurkanapithecus also widen somewhat superiorly, but are lon-ger, and have a different shape at the nasofrontal suture. Theexternal surface of the nasals is sellar, being slightly concavesuperoinferiorly and convex mediolaterally in lateral view.This morphology is found elsewhere only in stem catarrhines,cercopithecoids (including Victoriapithecus), and living plat-yrrhines, whereas the nasal bridge is flattened in living apes(although gorillas bear an autapomorphic stenotic condition),as well as all proconsuloids in which the region is preserved(Rae, 1999). The region is unfortunately not preserved inEpipliopithecus (Zapfe, 1960) and is badly distorted inLaccopithecus.

The roof of the nasal cavity preserves the superiormost re-mains of the midline nasal septum, which divides the narrowinterorbital portion of the cavity (Fig. 1b). On either side ofthis division, the remnants of the nasoturbinal ridges markthe boundary between the nasals and maxilla. The ascendingwing of the premaxilla on the right side makes contact witha substantial length of the nasal bone, leaving only the superiorhalf of the nasal in contact with the frontal process of the max-illa (Fig. 1a). This configuration is found in living platyrrhinesand stem catarrhines (e.g., Aegyptopithecus), and contrastswith the two extremes exhibited by living catarrhines (see

Fig. 2. Anterior views of Hylobates (a), Aotus (b), Cercopithecus (c), and Cebus

(d). Arrows in bed indicate infraorbital foramina. Arrow in (a) indicates

patent seam left behind by inferiorly drifting infraorbital canal (see text for

discussion).

Figure 1 in Rae, 1999). The short premaxilla of hominoidsis nearly excluded from contacting the nasal by the interveningmaxilla in most species (gorillas have the most overlap, but itis still considerably less than in non-hominoids). In extant cer-copithecoids premaxillary-nasal contact is far more extensive,with the premaxilla sometimes reaching the frontal (Rae,1999). The extent of this overlap in Victoriapithecus is ambig-uous at present (Benefit and McCrossin, 1993).

As mentioned above, the right-side break joins the interor-bital fragment with the lower face, making extensive contactbetween the premaxillary and maxillary fragments, includingthe lacrimal crest. The nasofrontal suture slopes inferolaterallyto either side of the midline to reach the maxilla. The contactbetween the frontal bone and the nasals and maxilla is an un-dulating, pneumatized surface like that of platyrrhines thatpossess a frontal sinus (Fig. 1b). The homology of the sinusin Lomorupithecus cannot be determined with certainty, butsince Aegyptopithecus, Proconsul, and Morotopithecus possessethmofrontal sinuses (Rossie, 2005), it is likely that the sinusin Lomorupithecus is the same. Regardless, it appears to bea considerably more inflated frontal sinus than is found inAegyptopithecus and is the first such occurrence in a small-bodied African catarrhine. Frontal pneumatization is also foundin Anapithecus (Kordos and Begun, 2000) and Epipliopithecus(Zapfe, 1960) but is of uncertain homology.

Nasal cavity. The right side lacrimal fossa is well preservedinternally and runs vertically to open into the inferior meatusof the nasal cavity, at which point there is a lateral hollowing,as in Kalepithecus (KNM-SO 417). The nasal aperture reachesbetween the orbits superiorly, which is common in anthropoidswith a wide interorbital region, such as colobines and gibbons.The aperture border is sharp throughout its length unlike mostother proconsuloids, in which it is more rounded inferiorly(Harrison, 1982). The V-shaped inferior border extends deeplybetween the incisor roots, as in many primitive catarrhines(e.g., Victoriapithecus, Aegyptopithecus), but unlike the condi-tion in Kalepithecus and Proconsul (Harrison, 1982). No atrio-turbinal ridge (see Rossie, 2005) is visible on the left side, andon the right side, the cortical bone is partly missing from theinternal face of the ascending wing of the premaxilla.

The floor of the nasal cavity is inferior to the level of thenasal aperture and nasoalveolar clivus such that a ‘‘stepped’’subnasal morphology is present (Ward and Kimbel, 1983).Portions of the palatine process of the maxilla are brokenaway, enlarging the incisive foramen, but as in most earlyMiocene catarrhines, the incisive canal appears to have beenlarge, basinlike rather than tubelike, and not roofed in the mid-line dorsally by the nasoalveolar clivus. The posterior borderof the foramen (or fenestra) was approximately in the mid-canine coronal plane. The alveolar part of the premaxilla iscompletely filled by the incisor roots except for the minutemissing portion near the midline. Maxillary sinuses invade themaxillary alveolar processes between the roots of M1 andreach the level of P4 anteriorly, as in many non-cercopithecoidcatarrhines (Ward and Pilbeam, 1983; Rossie, 2005). With theinterorbital fragment attached, the nasal aperture seemssomewhat narrow relative to its height, as in Dendropithecus,


and much more vertically oriented than in Victoriapithecus,Aegyptopithecus, and Afropithecus.

Palate. Small foramina are present bilaterally in the pre-maxillary portion of the palatal roof, anterior to the jaggedpremaxillomaxillary suture. These foramina are also presentin Epipliopithecus (Zapfe, 1960), Pliopithecus zhanxiangi(Harrison et al., 1991), and Micropithecus clarki (Fleagle,1975). The palate is shallow anteriorly, but deepens posteri-orly. Palatomaxillary sutures are present, reaching the midlineonly as far forward as the distal end of M1. They are morecoronally aligned than in M. clarki (Fleagle, 1975). The sutureon the right side runs into the trough for the greater palatineforamen. The toothrows diverge posteriorly, slightly morethan in M. clarki (Fleagle, 1975), but less than in more plesio-morphic catarrhines like Aegyptopithecus (Simons, 1987).

External snout. In lateral view the snout is very short, andthe plane of the nasal aperture is nearly vertical (Fig. 1).The canine jugum is large, with a slight fossa on the premax-illa anteriorly and a pronounced postcanine fossa that occupieswhat little space exists between the canine jugum and theorbital rim and malar region. The zygomatic processes ofthe maxilla are broken on both sides, but their remains clearlydemonstrate that the malar surface sloped posteroinferiorlyaway from the orbital rim. The junction of the malar surfacewith the body of the maxilla runs posteroinferiorly alonga straight line, in a nearly sagittal plane, as in gibbons, somecolobines, Aotus, Micropithecus clarki, Epipliopithecus, andLaccopithecus (Figs. 1 and 3). This contrasts with the more

Fig. 3. Lateral views of Macaca (a), Cebus (b), Aotus (c), and Hylobates (d).

Black line in each image traces the junction between the malar surface and the

body of the maxilla. Compare with Fig. 1d.

typical anthropoid conditions (Fig. 3a,b), in which this junc-tion occurs either along a curved, anteriorly convex line(e.g., Victoriapithecus, Cebus) or along a more vertical linewith a more coronal malar surface (e.g., Afropithecus, Aegyp-topithecus, Proconsul heseloni, Macaca). As in pliopithecoidsand gibbons, the malar surface terminates superomedially atthe infraorbital foramen in a plane posterior to that of theorbit rim, rather than anterior to it. On the right side, theanteroinferiorly facing part of the zygomatic process rootappears to have flared abruptly laterally, rather than sweepingposterolaterally, just inferior to the infraorbital foramen. Thebase of the zygomatic process root would have been at mostca. 3 mm above the alveolar margin. Such a low positionof the root contrasts with the higher root in gibbons, but iscommon among more plesiomorphic catarrhines like Epiplio-pithecus, Victoriapithecus, and Micropithecus clarki.

The malar region and midface of the specimen are rela-tively slightly taller than in M. clarki, judging from the posi-tion of the orbit. The base of the zygomatic process rootwould have been positioned above M1, and with the anteriorlacrimal crest lying above the P3/P4 junction, yields an evenmore orthognathic snout than in M. clarki. Most of the frontalprocess of the maxilla is preserved on the right side, and thesharp anterior lacrimal crest marks the inferomedial borderof the orbit, which lies anterior to the level of the infraorbitalforamen, as in M. clarki, gibbons, and pliopithecoids (Zapfe,1960; Fleagle, 1975).

Dentition. The upper central incisor roots are slightly widerthan those of the lateral incisors, to which they are positionedanteromedially (Fig. 1c). The roots are somewhat procumbent,but the crowns need not have been. A diastema separates I2

from the canine on the left side, and the premaxillomaxillarysuture is clearly visible within this space. The right canine isdisplaced laterally, causing breakage of the bone overlyingits jugum, and the medial portion of its alveolus is filledwith matrix, indicating that the damage was postmortem.

The left canine is represented by the root and base of thecrown, preserving the lingual cingulum, which starts at thedistal crown margin and rises minimally as it wraps aroundthe crown mesially. Distolingually, a wear facet for the lowerpremolar is present, as in Kalepithecus (KNM-SO 417) andDendropithecus (KNM-RU 1850), implying the presence ofa tall P3 protoconid. An unassigned P3 from Napak IV(UMP 66-06) has a tall enough protoconid to have producedthe wear facet on this canine and is as large as the teeth inthe BUMP 266 palate. It may belong to Lomorupithecus,but we refrain from making an attribution until associatedlower premolars are known for this species (see Appendix 1for discussion of pertinent previously collected material).The mesial part of the canine crown is broken away belowthe level at which the cingulum would have met the mesialgroove, the presence of which cannot be demonstrated. Inall preserved aspects, the tooth is most like that of Kalepithe-cus. The crown base is more buccolingually compressed thanis typical for pliopithecoids (except Epipliopithecus vindobo-nensis), but its lingual surface is less concave than inDendropithecus.


The P3 is mesiodistally short and buccolingually wide, witha triangular outline (Fig. 1c). It is the most mesiodistally com-pressed P3 of any early Miocene small-bodied catarrhine(Fig. 4; Table 1). Only Dendropithecus approaches this condi-tion. The distal fovea is much larger than the mesial fovea, withthe protocone narrower and more mesial than the paracone sothat the lingual margin of the crown angles distobuccallyfrom the protocone. This morphology is common in Dendropi-thecus and Aegyptopithecus and is approached by that of Lim-nopithecus evansi, but contrasts with the more evenly convexdistal margin of other Miocene small-bodied catarrhines. Thecrown is slightly worn, but the cusps are less heteromorphicthan in Kalepithecus and Dendropithecus. The morphology ofthe buccal face is much like Kenyan specimens of Micropithe-cus clarki (Leakey and Leakey, 1987)da rounded diamondshape with hints of buccal cingulum. The mesial transversecrest is distinct, but the distal crest is difficult to discern, partlydue to wear. The postprotocrista descends the protocone, encir-cling the distal crown margin. No lingual cingulum is presenton the bulbous protocone, which has a more flared lingualface than in Dendropithecus or Kalepithecus.

The P4 has a more symmetrical crown outline than P3 andmore evenly sized anterior and posterior foveae (Fig. 1). Thedistal cingulum ascends the protocone, producing a slight,rounded lingual cingulum, somewhat like Kalepithecus butnot as distinct as in Simiolus enjiessi (Leakey and Leakey,

Fig. 4. Relative breadth of upper third premolars. Box plot of mesiodistal

length/buccolingual breadth� 100 in Lomorupithecus (n¼ 1), Dendropithecus

macinnesi (n¼ 7), Limnopithecus evansi (n¼ 4), Simiolus enjiessi (n¼ 1),

Limnopithecus legetet (n¼ 3), and Micropithecus clarki (n¼ 3). Boxes delimit

the 25th and 75th percentiles. Whiskers delimit 5th and 95th percentiles. Data

are from Harrison (1982) and Leakey and Leakey (1987).

Table 1

Dental metrics (mm) of BUMP 266 (left side) and BUMP 269

BUMP 266 BUMP 268

C P3 P4 M1 M1 M2

MD 6.2 3.5 3.6 4.9 5.2 6.2

BL 4.9 6.0 6.2 6.2 4.6 5.2

1987). The buccal surface of the paracone has a more lingualcant than in Simiolus, resembling Limnopithecus legetet andKalepithecus. Both transverse cristae were present, as evi-denced by a distinct central fovea. Small pre- and postpara-styles are present instead of a continuous buccal cingulum.

The left M1 is quite worn, to just above the plane of the lin-gual cingulum, which is pronounced lingually to the protoconeand continues around the hypocone to end at the distobuccalcorner of the crown. The metacone and hypocone are all butworn away, but their bases can be distinguished from the cin-gulum. A small ledge of cingulum is present buccally betweenthe paracone and metacone. The tooth is short and broad, witha rounded distolingual margin and a somewhat distallypositioned hypocone.

The very worn right M1 preserves a mesiodistally shortdistal fovea and shows that the talon was small compared to thetrigon, as in Micropithecus clarki and Limnopithecus evansi.Unlike M. clarki, the M1 crowns are mesiodistally short andhave a more prominent cingulum at the mesiolingual corner.The broken and worn crowns of right P3eM1 add little infor-mation, but the M1 confirms that the lingual cingulum is quitebroad lingual to the protocone, in contrast to the condition inM. clarki. Such a pronounced lingual cingulum is also presentin Epipliopithecus, Pliopithecus, Platodontopithecus, andDionysopithecus (Begun, 2002).

BUMP 268

Specimen BUMP 268 is the left half of a juvenile mandiblefrom Napak IX preserving alveoli for I1, I2, dc, and dm1, rootsand a partial crown of dm2, and crowns of M1 and M2 (Fig. 5).The M2 is unerupted but partly exposed and was further ex-posed during preparation by the removal of overlying alveolarbone. Radiographs of the hemi-mandible reveal the presenceof unerupted premolar crowns below the deciduous molarroots. The occlusal area (length� breadth; see Table 1) ofthe M1 in BUMP 268 is similar to that of Limnopithecus evansiand Micropithecus clarki, but the crown is broader, its breadth/length ratio falling within the range of the larger Kalepithecus,Aegyptopithecus, and Epipliopithecus vindobonensis, as wellas the much larger Dendropithecus (Fig. 6b). Since the M1

of BUMP 266 is also within the M. clarki size range (in termsof occlusal area), but is buccolingually broader, both speci-mens most likely belong to Lomorupithecus (Fig. 6a). Theonly other taxa positively identified from Napak are the largerDendropithecus, Micropithecus clarki, which is smaller andhas narrower molar crowns, and Limnopithecus legetet, whichhas larger and narrower crowns with sharper cusps and cristids(Harrison, 1982). The M1 of BUMP 268 is similar to those ofL. evansi in proportions (Fig. 6b) but not in morphology (seebelow).

The corpus is shallow and gracile, unlike juvenile mandi-bles of Limnopithecus legetet (Harrison, 1982), and increasesslightly in depth anteriorly. The planum alveolare slopessteeply down to the superior transverse torus, below whichthere is a small but distinct digastric fossa. There is no inferior


Fig. 5. Juvenile mandible of Lomorupithecus harrisoni BUMP 268: medial (a) and occlusal (b) views of medial prior to removal of bone overlying M2, (c) enlarged

view of M1, (d) illustration of M1, and (e) enlarged view of M2 after removal of overlying bone. Scale bar is for a and b only.

transverse torus. A long but shallow sub-mandibular fossa runsfrom under the M1 level posterosuperiorly to end at the distalbreak, at which point the groove for the mylohyoid nerve ispresent. What remains of the dm2 crown reveals only that itwas much narrower mesially than distally.

The M1 (Fig. 5c,d) is slightly narrower in its mesial moi-ety (4.1 mm) than in its distal moiety (4.6 mm). The enamelof the buccal cusps appears to have been etcheddeither byacidic soil conditions or by the digestive acids of some pred-ator. All of the cusps are low and blunt, and the cristids arerounded and poorly defined. A narrow buccal cingulumruns from the preprotocristid to the hypoconulid, interruptedonly at the distobuccal face of the hypoconid. The protoconidis lingual to the buccal crown margin, causing the cristid ob-liqua to run forward to the protoconid at a slightly obliqueangle. The metaconid is distal to the protoconid, giving thecusps a staggered appearance, as in Micropithecus clarki,Kalepithecus, Limnopithecus evansi, and pliopithecoids. The

trigonid, or mesial fovea, is relatively large and longer thanin other small-bodied proconsuloids. Unlike M. clarki, thehypoconulid is very small and placed buccal to the midline ofthe crown, leaving room for a larger distal fovea, which fillsthe area between the hypoconulid and entoconid. Comparedto Limnopithecus legetet and M. clarki, the M1 is broader,has a larger distal fovea, and a more closely appressed hypo-conid and hypoconulid. Compared to the new specimen, L.evansi has more distally and medially placed hypoconulidsand less mesial tapering of the crown. Harrison (1982) notedthat some of the unassigned lower molar material from Napakresembled L. evansi metrically and morphologically, andwhile the crown dimensions of BUMP 268 M1 are similarto a large specimen of L. evansi, its morphology (and thatof the M2) differentiates it from that taxon, which remainsknown only from Songhor.

The cusps of the M2 are more peripherally placed on theocclusal surface than in the M1, and the talonid basin and


distal fovea are very deep and wide (Figs. 5e, 7a). The lingualcusps are quite narrow buccolingually, and the cristids con-necting them are sharp and prominent. In contrast, the buccalcusps are more inflated and crowded together. The cristid ob-liqua is short, and the hypoconulid is appressed tightly againstthe hypoconid such that the interposing cristids are barely dis-cernible. A distinct cristid descends into the talonid from theapex of the hypoconid in a position that is identical to thatof the distal arm of the ‘‘pliopithecine triangle’’ of pliopithe-coids. The distal fovea is very large, being both broad(w50% of crown width) and long (almost a third of crownlength). Its great length is produced by three factors: 1) theplacement of the entoconid, which is nearly as mesial as thehypoconid; 2) the mesial crowding of the hypoconulid against

Fig. 6. Scatter plots of M1 (above) and M1 (below) crown proportions (Y-axis)

and occlusal area (X-axis) for a comparative sample of small catarrhines.

Combination of crown proportions and crown size provides a basic illustration

of the ‘‘morphospace’’ occupied by various taxa. Units are in millimeters. Data

are from Zapfe (1960), Kay et al. (1981), Harrison (1982), Leakey and Leakey

(1987), and Harrison and Gu (1999).

the hypoconid; and 3) the mesial position of the postcristid,which runs buccally from the most mesial portion of the hypo-conulid. The postentocristid is accordingly long, and arcs dis-tobuccally around the fovea.

The mesial fovea is mesiodistally shorter than in M1 butstill long compared to the M2 of other small-bodied proconsu-loids. The hypometacristid and hypoprotocristid that form itsdistal border are distinct and slightly oblique to the longaxis of the crown, as in Micropithecus clarki, Simiolus enjiessi,Dendropithecus, and pliopithecoids. The long paracristid runsmesially toward the crown margin before turning lingually ata sharp angle to enclose the fovea mesially. This configurationproduces a mesially protruding prowlike mesiobuccal cornerto the fovea, which is not common in other small-bodiedproconsuloids (Fig. 7) but appears in Dionysopithecus, Plato-dontopithecus, Anapithecus (Begun, 1989), and Pliopithecus(Begun, 2002). The twinning of the hypoconid and hypoco-nulid contrasts with the more median hypoconulid seen in pro-pliopithecoids and proconsuloids but is similar to the conditionin some pliopithecoids (e.g., Platodontopithecus and Pliopithe-cus antiquus).

In sum, the combination of features in the lower M1 and M2

of BUMP 268 distinguish it from Dendropithecus, Micropithe-cus, Kalepithecus, and Limnopithecus. The mesially taperingM1 crown bearing a slightly oblique cristid obliqua and stag-gered mesial cusps, combined with a non-tapering M2 bearinga straighter cristid obliqua, a large and distally protrudingdistal fovea, and a weak buccal cingulum are similar to con-ditions found in Platodontopithecus. The size and morphologyof the mesial fovea and paracristid of both molars are moresimilar to morphology observed in Pliopithecus platyodonand Pliopithecus antiquus. Overall, the morphology of theLomorupithecus lower molars is consistent with that of a basalpliopithecoid.

The facial morphology of Lomorupithecus also resemblesthat of pliopithecoids in many respects, but many featuresare also shared with gibbons, and a few are shared with Micro-pithecus clarki. The similarities to pliopithecoids and gibbonsmay be symplesiomorphies, but recent studies suggest that thefacial features shared by pliopithecoids, gibbons, and colo-bines are not primitive for catarrhines (Benefit and McCrossin,1991, 1993, 1997). The uncertain polarity of these featuresmakes their relevance to the phylogenetic position of Lomor-upithecus unclear. Accordingly, in order to investigate thephylogenetic position of the new species, a cladistic analysisof 36 extant and fossil anthropoids was conducted.

Phylogenetic analysis

Coding of the 191 craniodental and postcranial characters(Appendix 2) was based on study of original material in theNational Museum of Kenya, Uganda Museum of Paleontol-ogy, Yale Peabody Museum, and Carnegie Museum of NaturalHistory, as well as published descriptions. Some of the charac-ters employed were delineated quantitatively, but many othersare qualitative (see Appendix 2). While we are acutely aware


Fig. 7. Second lower molars of (a) Lomorupithecus harrisoni BUMP 268, (b) Limnopithecus legetet KNM-CA 1810, (c) Micropithecus clarki KNM-CA 380, (d)

Pliopithecus antiquus (type specimen), (e) Limnopithecus evansi KNM-SO 444, (f) Kalepithecus songhorensis KNM-SO 378; b, c, e, and f are modified from

Leakey and Leakey (1987). Scale bar¼ 1 mm.

that the compilation of morphological character matrices isplagued with pitfalls, ranging from the possibility of correlatedcharacters to subjective character state definition (e.g., Pogueand Mickevich, 1990; Gift and Stevens, 1997), we feel thatthe benefits of presenting character data in this explicit manneroutweigh the drawbacks. Two unweighted analyses were per-formed: one with all characters unordered, and one with someof the characters ordered (see Appendix 2) to reflect hypothe-sized character-state transformations (Slowinski, 1993; Wiens,2001).

Heuristic searches in PAUP Version 4 (Swofford, 1993)using 10,000 replicates yielded 18 equally most-parsimonioustrees (860 steps in length) for the ordered analysis and six(814 steps in length) for the unordered analysis. The strictconsensus trees for each analysis place Lomorupithecus inthe Eurasian pliopithecoid clade (Figs. 8, 9a). In the orderedanalysis, Lomorupithecus is nested within Pliopithecoidea,whereas in the unordered analysis it is placed at the base ofthis clade. Apart from the pliopithecoid clade, both analysesrecovered the clades Nyanzapithecinae (see Harrison, 2002),Cercopithecoidea, and Hominoidea, but other relationshipsare ambiguous. Since we support the ordering of certain char-acters, we favor the results of the ordered analysis (Fig. 8), butthe basal position of Lomorupithecus recovered by the un-ordered analysis seems more biogeographically plausible. Aconsensus of all 24 trees from both analyses (Fig. 9b) shows

the points of agreement between the two, which includea Lomorupithecus-pliopithecoid clade.

Character tracing on the results of the ordered analysisindicates that the position of Lomorupithecus is supported by17 synapomorphies shared with other pliopithecoids (nine are un-ambiguous; Table 2). Many of these are local synapomorphiesthat are also found elsewhere among catarrhines (e.g., meta-cone size, orthognathic snout), but a few are more narrowlydistributed. In particular, the narrow M1e2 mesial fovea witha prowlike paracristid, narrow and distally projecting distalfovea, and twinned hypoconulid and hypoconid are foundonly in Lomorupithecus and some pliopithecoids (Fig. 7).Pliopithecoidea is usually defined by a suite of distinctive den-tal features, including waisted upper and lower incisors, tall P3

protoconid with a short mesial face, ‘‘pliopithecine triangle’’on lower molars, staggered mesial cusps and oblique cristidobliqua on M1e2, mesially narrow M1 crowns, and broad up-per molars with a broad lingual cingulum (Andrews et al.,1996; Harrison and Gu, 1999; Begun, 2002). No P3s or inci-sors of Lomorupithecus are yet known, but with the exceptionof the ‘‘pliopithecine triangle,’’ the remaining features arepresent in Lomorupithecus, and thus it is not surprising thatit emerges as a pliopithecoid in this analysis. It is noteworthythat these molar features are variably expressed among pliopi-thecoid species, and that the ‘‘pliopithecine triangle’’ is alsonot present in Laccopithecus (and most other crouzeliins)


and is variably absent in Epipliopithecus (Begun, 2002).Moreover, Lomorupithecus exhibits a particular combinationof states found only in pliopithecoids. The overall gibbonlikefacial morphology (characters 78, 82, 88, 94 in Lomorupithe-cus) could link Lomorupithecus with gibbons or even colo-bines, but when combined with the primitive catarrhineconfiguration of the premaxilla and nasal bones, it is most par-simoniously placed among the pliopithecoids. The combinedeffect of the dental and cranial similarities to pliopithecoidsis a strong phylogenetic signal, despite the absence of someof the ‘‘classic’’ pliopithecoid features.

The bootstrap value for the pliopithecid clade, includingLomorupithecus, is 63% (1000 replicates), and the relationshipis not affected by experimental removal of cercopithecoids,nyanzapithecines, or the other African small catarrhines (i.e.,it is not an artifact of sampling bias). Accordingly, we regardthis phylogenetic hypothesis as well-supported. However,given the fragmentary nature of so many of the Miocene

Fig. 8. Results of ordered cladistic analysis. Strict consensus tree of 18 equally

parsimonious cladograms (860 steps) for 36 taxa and 191 morphological char-

acters. CI¼ 0.32, RI¼ 0.60, RCI¼ 0.19. Bootstrap values (1000 replicates)

are: node 1 (Catarrhini) 96%, node 2 (Pliopithecidae) 63%, node 3 (Pliopithe-

cinae) 49%, node 4 (Cercopithecoidea) 100%, node 5 (Nyanzapithecinae)

70%, node 6 (Hominoidea) 78%.

catarrhines included in our analysis, we recognize that betterknowledge of their anatomy could alter this phylogenetic pic-ture significantly. In particular, if many of the other AfricanMiocene taxa are found to have pliopithecoid-like cranialanatomy combined with unambiguous crown-catarrhine traits(e.g., lost entepicondylar foramen, tubular ectotympanic),then it would likely demonstrate that the facial similaritiesbetween Lomorupithecus and pliopithecoids are symplesio-morphic. In such an event, the relationship between the twocould very well break down. Considering the important pale-obiogeographical implications of our phylogenetic results,the phylogenetic position of Lomorupithecus should be reeval-uated if such material comes to light.

The precise position of Lomorupithecusdeither as a pliopi-thecin (Fig. 8) or a stem pliopithecoid (Fig. 9a)dis ambi-guous in our analysis. Its inclusion in the pliopithecin tribein the ordered analysis is based largely on its very broad M1

crowns and the twinned hypoconid and hypoconulid on thelower molars (Table 2). The former is a local synapomorphysince it represents a reversal to a primitive catarrhine condi-tion (Laccopithecus and Dionysopithecus have slightly nar-rower crowns), and thus the state in Lomorupithecus wouldbe equally appropriate for a stem pliopithecoid. The twinnedhypoconid and hypoconulid appears to be of less ambiguouspolarity, but it is impossible to say whether a slight increasein homoplasy in this character is less likely than the implau-sible biogeographic scenario that its optimization wouldrequire.

The basal position of Lomorupithecus favored by the unor-dered analysis is less problematic biogeographically, but manyof the precise relationships recovered by that assumption setare dubious. The sister-taxon relationship of Dionysopithecusand Pliopithecus runs counter to current views (Harrison andGu, 1999), and the positions of Catopithecus and Aegyptopi-thecus are certainly erroneous. Given the conflict betweenthe results of the two assumption sets, it may be best to con-sider the combined consensus tree (Fig. 9b) as representativeof the level of precision that can currently be achieved forthe Miocene taxa. In view of this uncertainty, we classifyLomorupithecus as incertae sedis at the family level.

Discussion

The phylogenetic position of the small-bodied catarrhinesfrom the Miocene of East Africa is far from clear, and al-though they are often treated as a natural group, it is certainlypossible that they constitute a paraphyletic or polyphyleticassemblage (e.g., Andrews, 1978; Harrison, 1988). Most recentauthors have expressed doubt that any of these species arestem hominoids but have refrained from offering more precisehypotheses on the basis of the limited material available (e.g.,Rose et al., 1992; Harrison, 2002). Since the group shares lit-tle more than their diminutive size in common, their relation-ships will certainly become clearer if and when sufficientcranial and postcranial material can be recovered. Indeed,the recently proposed clade Dendropithecoidea (Dendropithecus,


Fig. 9. Results of unordered cladistic analysis. (a) Strict consensus of six trees from unordered analysis (814) steps. CI¼ 0.33, RI¼ 0.58, RCI¼ 0.19. (b) Strict

consensus of combined ordered and unordered trees showing points of agreement between the two assumption sets. Note that the unordered analysis places

Lomorupithecus in a basal position within the pliopithecoid clade. See text for discussion.

Simiolus, and Micropithecus; Harrison, 2002) is largely basedon postcranial similarities gleaned from recent discoveries(Rose et al., 1992). But considering how little is known aboutthe anatomy of most of these small catarrhines, it is probablybest not to assume a great deal about their true phylogeneticaffinities. For instance, the stem cercopithecoid and stemhominoid lineages may stretch into the middle if not earlyOligocene (Steiper et al., 2004), and it is certainly possible thatcercopithecoids more primitive in appearance than victoriapi-thecids survived until the early Miocene. Such taxa would bedifficult to discriminate from stem catarrhines, particularly onthe basis of limited dental material. Similarly, the resemblancebetween plesiomorphic pliopithecoids from Asia and the Afri-can proconsuloids and dendropithecoids led Harrison and Gu(1999) to comment that an earlier pliopithecoid (than those atSihong) may already be known and ‘‘lurking undetectedamong the poorly known smaller proconsulids from EastAfrica’’ (p. 268). They doubted that this was the case at thetime, and cautioned that if such a species were to be discov-ered, its pliopithecoid affinities could be very subtle. Weagree with this assessment, and it is because of the subtletyand uncertain polarity of the similarities between Lomorupi-thecus and pliopithecoids that we undertook a cladistic anal-ysis of the new species along with a broad array of living and

extinct taxa in order to determine their phylogenetic signifi-cance. The possibility that Lomorupithecus is a pliopithecoidmay seem surprising, but it may be an indication of the kindof taxonomic diversity that will be revealed when other small-bodied Miocene catarrhines are known from more completematerial.

If Lomorupithecus is a pliopithecoid, then it is the firstmember of this group to be found on the Afro-Arabian land-mass (although some intriguing pliopithecoid-like traits havebeen noted in a mandibular specimen from Fort Ternan thatis usually attributed to Limnopithecus; Begun, 2002). It isolder than the oldest Eurasian pliopithecoids (Harrison andGu, 1999), which might suggest that it is close to the ancestryof the Eurasian clade, but without knowing its precise phylo-genetic position within Pliopithecoidea, its biogeographicalsignificance is ambiguous. The implications of its differentplacements in the ordered and unordered analyses are quitedissimilardthe former requiring either multiple dispersalsout of Africa or a return to Africa from Eurasia by the lineageleading to Lomorupithecus. Obviously, a basal position forLomorupithecus is more biogeographically parsimonious.

The morphology of Lomorupithecus may have implicationsfor hypotheses regarding the polarity of cranial featuresamong major catarrhine clades. In the traditional view, the


features shared by living gibbons, colobines, and the fossilpliopithecoids are believed to reflect the primitive conditionfor crown catarrhines (e.g., Vogel, 1968; Delson and Andrews,1975; Harrison, 1982, 1987). More recently, Benefit andMcCrossin (1991, 1993, 1997) have proposed that the featuresshared by Aegyptopithecus, Afropithecus, and Victoriapithecusrepresent the ancestral catarrhine condition. The latter hypoth-esis clearly owes its inception to the discovery of well-preserved cranial specimens belonging to Neogene catarrhines,and further recovery of such fossils will be critical in testingboth hypotheses. Character tracing on our strict consensustree yields an ancestral crown catarrhine morphotype (thebranch just below the cercopithecoid-hominoid split) that iscongruent with that hypothesized by Benefit and McCrossin(e.g., 1993), in including a non-protruding orbital rim and broadinterorbital region, but it differs in having orbits positionedlow on the face and a posteroinferiorly sloping malar surface.Unfortunately, several facial characters are ambiguous on thisbranch, including nasal length, snout length, presence ofa frontal trigon, and presence of supraorbital costae or tori.Consensus trees are not the best frameworks on which to tracecharacter evolution (Barrett et al., 1991), and our results arecertainly compromised by the large polytomy that subsumesall of the potential crown catarrhines. Obviously, these resultsare also affected by the pliopithecoid placement of Lomorupi-thecus in our analysis. Had Lomorupithecus taken a positionalong the catarrhine stem lineage, it might have had a greater

Table 2

Character support for nodes within Pliopithecoidea observable in

Lomorupithecus

Character CI Derived character state

Pliopithecidae (node 2)

36 0.273 Hypocone more lingual

than protocone*

54 0.500 M1 crown tapering anteriorly*

55 0.400 M1 cristid obliqua slightly oblique

56 0.273 M2 crown narrow*

58 0.333 M1e2 paracristid prowlike

65 0.400 M2 cristid obliqua oblique (intermediate)

78 0.500 Orbit inferior margin barlike and protruding

82 0.200 Nasal bones short

88 0.143 Snout orthognathic

94 0.200 Junction of malar surface and

body of maxilla slopes posteroinferiorly

Pliopithecinae (node 3)

18 0.154 P4 crown ovoid*

55 0.400 M1 cristid obliqua oblique

(reverts to slightly oblique)

62 0.444 M1e2 hypoconulid buccal

(twinned in Lomorupithecus)*

69 0.250 M1e3 distal fovea narrow and projecting*

Pliopithecini

21 0.273 M1 crown very broad*

31 0.111 Metacone size� protocone*

62 0.444 M1e2 hypoconulid twinned with hypoconid*

Parentheses after state change indicate state in Lomorupithecus, CI¼consistency index. Characters traced on results of the ordered analysis (Fig. 8).

*Indicates unambiguous transition.

effect on global polarity, but as a pliopithecoid, its anatomysimply conforms to that of its close relatives. While this resultin itself implicitly supports the revised view of the catarrhineancestral morphotype (e.g., Benefit and McCrossin, 1993), theambiguity of relationships among potential crown catarrhinesin our analysis prevents a more meaningful test of thehypothesis.

Conclusions

Lomorupithecus harrisoni is known, in part, from one of themost complete facial specimens of an African small-bodiedMiocene catarrhine. In many ways, its facial and dentalmorphology resembles that of the Eurasian pliopithecoids.Our cladistic analysisdintended to explore the phylogeneticaffinities of the new taxondindicates that it is a member ofPliopithecoidea. As such, it would be the oldest knownpliopithecoid and the only member of that clade to be foundon the Afro-Arabian landmass. As with any phylogenetichypothesis, this result will be tested by each new catarrhinefossil that comes to light, but at present it is the most parsimo-nious explanation of the available data.

Acknowledgments

We thank William Downs, Moses Mafabi, Ezra Musiime,and Robert Kityo for their work in the field. We are gratefulto the Uganda National Council for Science and Technologyand Dr. Epraim Kamuhangire, Commissioner of Antiquities,for permission to undertake research in Uganda. We thankEric Seiffert, David Begun, Terry Harrison, and two anony-mous reviewers for helpful discussions and comments onthis manuscript, Eric Sargis and David Pilbeam for accessto the cast collections in their care, and Meave Leakey,Mary Muungu, and Christopher Kiarie for access to fossilsin the National Museum of Kenya. This research was sup-ported by grants from the LSB Leakey Foundation, theWenner-Gren Foundation, and the National Science Founda-tion (BCS-0456589) to L.M. and a Rea Postdoctoral Fellow-ship to J.R.

Appendix 1. Previously collected specimens from Napak

The Napak sequence has previously produced craniodentalmaterial attributed to Micropithecus clarki (Fleagle, 1975;Fleagle and Simons, 1978), Dendropithecus macinnesi (Har-rison, 1982), and Limnopithecus legetet (Fleagle and Simons,1978), but a number of previously collected specimens fromthese localities (primarily lower anterior teeth) indicated thepresence of an unidentified fourth small-bodied non-cercopi-thecoid catarrhine (Harrison, 1982). The specimens describedhere confirm the presence of a fourth small-bodied catarrhine,but we refrain from attributing any of the previously unas-signed specimens to Lomorupithecus until the anterior lowerdentition of the taxon is known. Only two other primate


specimens have been collected from Napak IX in the past:a frontal bone, and a mandible with two worn teeth, neitherof which can be confidently attributed to Lomorupithecus.

Specimen UMP 66-23, a mandibular fragment with M2e3,is too heavily worn to reveal any details of occlusal anatomy.The size and outline of the molars are compatible with attri-bution to M. clarki, and the M3 appears to have the reducedentoconid typical of that species. Harrison (1982) retainedthis specimen in M. clarki, and we see no reason to questionthis attribution. The frontal bone UMP 68-25 was originallyattributed to an unnamed cercopithecoid (Pilbeam andWalker, 1968), but it was later included in the hypodigm ofM. clarki (Fleagle and Simons, 1978). The interorbital regionof the specimen is considerably narrower than that of Lomor-upithecus, and the angle formed by the orbital roof and inter-orbital region in the sagittal plane indicates a more prognathicmidface. The sharp margin and acute angle between the or-bital roofs and frontal squama are unlike the condition seenin non-cercopithecoid catarrhines, and it is our opinion thatthe specimen most closely resembles the living cercopithe-coids. Of course, its attribution to Micropithecus cannot beentirely ruled out until the upper face of that genus is betterknown.

Appendix 2. Character descriptions and matrix

Ordered characters: 5, 7, 9, 11, 16e19, 21, 22, 26, 27, 29,34, 35, 38, 39, 47, 50, 53, 55e57, 62, 63, 66, 70, 72, 80, 84,97e99, 102, 107, 108, 110, 120, 122, 127, 130, 132, 134, 135,137, 149, 150, 161, 163, 167, 174, 184.

Character descriptions

1. I1 shape: (0) labiolingually thin; (1) labiolingually thick.2. I2 size: (0) much smaller than I1; (1) slightly smaller than

I1.3. I2 cingulum: (0) present; (1) absent.4. Incisor crown waisting: (0) absent; (1) present.5. Upper canine bilateral compression: (0) very compressed;

(1) compressed; (2) less compressed.6. P2: (0) present; (1) absent.7. Upper premolar cusp heteromorphy: (0) marked; (1)

reduced; (2) subequal.8. Upper premolar cusp volume: (0) buccolingually narrow;

(1) inflated, not crowded; (2) inflated and crowded.9. P3 crown shape: (0) triangular; (1) sub-ovoid; (2) very

ovoid.10. P3 distal margin: (0) convex; (1) angled; (2) straight.11. P3 proportions: (0) very broad; (1) broad; (2) narrow; (3)

very narrow (mesiodistally expanded).12. P3 transverse crest: (0) links cusps; (1) cusps separated

by groove.13. P3 buccal cingulum: (0) absent; (1) present.14. P3 protocone lingual face: (0) sloping; (1) vertical.15. P3 buccal face: (0) triangular; (1) diamond-shaped; (2)

with mesiobuccal expansion on root.

16. P3 lingual cingulum: (0) inflated; (1) present; (2) absent.17. P4 proportions: (0) very broad; (1) broad; (2) narrow; (3)

very narrow.18. P4 crown shape: (0) triangular; (1) sub-ovoid; (2) ovoid.19. P4 lingual cingulum: (0) inflated; (1) present; (2) absent.20. Molar cusp form: (0) high and sharp; (1) low and

rounded; (2) high and inflated; (3) high and very inflated.21. M1 crown: (0) very broad; (1) broad; (2) narrow; (3) very

narrow.22. M2 crown: (0) very broad; (1) broad; (2) narrow; (3) very

narrow.23. M1 metacone and hypocone size: (0) similar to mesial

cusps; (1) smaller than mesial cusps.24. M2 cusp at mesiolingual corner of crown: (0) absent; (1)

present.25. M1eM2 waisting: (0) absent; (1) present.26. M1eM2 crown tapering: (0) none; (1) distal tapering; (2)

strong distal tapering.27. M1eM3 lingual cingulum: (0) strong; (1) reduced; (2)

none.28. M2eM3 buccal cingulum: (0) present; (1) absent.29. M1eM2 mesial shelf: (0) none; (1) narrow cingulum; (2)

pronounced cingulum; (3) shelf.30. M3 distal moiety: (0) reduced; (1) unreduced.31. Metacone size: (0) <protocone; (1) �protocone.32. Protocone position: (0) aligned with paracone; (1) distal

to paracone and crown margin.33. Crista obliqua: (0) present; (1) absent.34. Protoconule size: (0) large; (1) small; (2) absent.35. Hypocone-metacone crista: (0) discontinuous; (1) contin-

uous; (2) true hypoloph.36. Hypocone position: (0) in line with metacone; (1) distal

to metacone; (2) distobuccal; (3) more lingual thanprotocone.

37. Prehypocrista position: (0) meets base of protocone; (1)meets crista obliqua.

38. Trigon size: (0) broad; (1) narrow; (2) very narrow andcrescentic.

39. Talon size: (0) absent; (1) small; (2) large relative totrigon.

40. M1eM3 size sequence: (0) M1<M2<M3; (1) M1<M3<M2; (2) M1wM3<M2; (3) M3<M1<M2; (4)M3<M2<M1.

41. I1 shape: (0) narrow and tall; (1) broader and shorter.42. I2 shape: (0) narrow and tall; (1) broader and shorter.43. I2 lingual cingulum: (0) absent; (1) present.44. Lower incisor lingual enamel: (0) present; (1) absent.45. I2 distal margin: (0) angled; (1) straight.46. P3 alignment: (0) oblique to tooth row; (1) nearly in

line.47. P3 sectoriality: (0) non-sectorial; (1) poor; (2) moderate;

(3) moderate to strong.48. P3 mesiolingual beak: (0) absent; (1) present.49. P3 metaconids: (0) present; (1) absent.50. P4 length: (0) very narrow; (1) narrow; (2) broad; (3) very

broad.51. P4 buccal flare: (0) slight; (1) strong.


52. P4 mesial fovea height: (0) slightly above distal basin; (1)much higher than distal basin.

53. M1 crown shape: (0) very narrow; (1) long, narrow; (2)broad; (3) very broad.

54. M1 crown tapering: (0) tapering anteriorly; (1) no tapering.55. M1 cristid obliqua: (0) oblique; (1) slightly oblique; (2)

straight to protoconid.56. M2 crown shape: (0) Very narrow; (1) long, narrow; (2)

broad; (3) very broad.57. Pliopithecine triangle: (0) none; (1) variable; (2) present.58. Paracristid: (0) curved; (1) beaklike.59. Protoconid-metaconid: (0) oblique; (1) transverse.60. M1eM2 mesial lophid: (0) absent; (1) present.61. M1eM2 paraconids: (0) present; (1) absent.62. M1eM2 hypoconulid: (0) absent; (1) appressed to

entoconid; (2) median; (3) buccal; (4) twinned withhypoconid.

63. M1eM2 crown outline: (0) rectangular; (1) slightly ovoid;(2) ovoid.

64. M1eM2 hypoconulid: (0) present; (1) absent.65. M2 cristid obliqua: (0) straight; (1) slightly oblique; (2)

oblique.66. M1eM3 buccal cingulum: (0) well-developed; (1) only in

notches; (2) none.67. M1eM3 lingual notch depth: (0) shallow; (1) deep open-

ing into talonid.68. M1eM3 mesial fovea: (0) broad; (1) narrow.69. M1eM3 distal fovea: (0) wide; (1) narrow; (2) narrow and

distally projecting.70. M3 shape: (0) relatively untapered; (1) tapers distally; (2)

tapers strongly, entoconid indistinct.71. M3 hypoconulid and entoconid linked by crest: (0) yes;

(1) no.72. M3 hypoconid and entoconid linked by crest: (0) no; (1)

yes; (2) true distal lophid.73. Inferior transverse torus: (0) present; (1) absent.74. Mandibular condyle height: (0) lower than coronoid; (1)

equal or higher.75. Superior transverse torus position: (0) at 1/3 height of

symphysis; (1) mid-symphysis height.76. Premaxilla ascending wing: (0) broad; (1) narrow.77. Clivus orientation: (0) protruding; (1) more vertical.78. Orbit inferior margin: (0) flush with cheek; (1) sharp,

protruding; (2) barlike, protruding.79. Orbit position: (0) high; (1) low.80. Pyriform inferior border: (0) narrow, V-shaped; (1) wider

and more rounded; (2) flat, horizontal.81. Nasal shape: (0) narrow superiorly; (1) broad superiorly.82. Nasal length: (0) long; (1) short.83. Nasal bridge: (0) flat; (1) rounded horizontally.84. Premaxillaenasal contact: (0) excluded or minimal; (1)

mid-nasal; (2) extensive.85. Atrioturbinal ridge: (0) present; (1) absent.86. Palate shape: (0) narrow anteriorly; (1) parallel tooth

rows; (2) bowed laterally.87. Horizontal palatine process: (0) broad; (1) narrow.88. Snout length: (0) protruding; (1) orthognathic.

89. Facial profile: (0) prosthion, rhinion, glabella in line; (1)stepped.

90. Buccal pouches: (0) absent; (1) present.91. Frontal sinus: (0) present; (1) absent.92. Infraorbital foramen position: (0) low; (1) very near

orbital rim.93. Interorbital region: (0) broad; (1) narrow.94. Infraorbital surface of maxilla: (0) slopes posteroinfer-

iorly; (1) slopes anteroinferiorly.95. Supraorbital region: (0) costae; (1) tori or arches; (2)

strong inflated tori.96. Frontal trigon: (0) present; (1) absent.97. Postglenoid foramen: (0) large; (1) reduced; (2) absent.98. Ectotympanic: (0) annular; (1) semi-tubular; (2) tubular.99. Lumbar centra: (0) dorsoventrally compressed and small;

(1) expanded; (2) very expanded.100. Vertebral body keeling: (0) present; (1) absent.101. Transverse processes: (0) on body; (1) on pedicle.102. Transverse process inclination: (0) ventrally inclined; (1)

neutral; (2) dorsally and caudally inclined.103. Accessory processes: (0) present; (1) absent or reduced.104. Spinous process inclination: (0) cranially inclined; (1)

caudally inclined.105. Lumbar centrum hollowing: (0) present; (1) absent or

reduced.106. Lumbar centra spooling: (0) present; (1) absent or

reduced.107. Lumbar number: (0) 7; (1) 6; (2) 5; (3) 4.108. Sacral number: (0) 3; (1) 4; (2) 5; (3) 6.109. Ischial spine: (0) distal to acetabulum; (1) level with

acetabulum.110. Acetabulum: (0) expanded with raised lip; (1) cranially

expanded lunate surface; (2) symmetrical lunate surface.111. Sacrum shape: (0) broad; (1) narrow.112. Scapula shape: (0) vertebral border< caudal; (1) long

vertebral border.113. Scapular spine: (0) oblique to axial border; (1) low angle

to axial border.114. Glenoid curvature: (0) more curved craniocaudally

than dorsoventrally; (1) uniformly and moderatelycurved.

115. Glenoid orientation: (0) faces laterally; (1) facescranially.

116. Scapula position: (0) lateral; (1) dorsal on thorax.117. Sternum: (0) long narrow; (1) short broad.118. Humeral head orientation: (0) <15 degrees; (1) >15

degrees.119. Dorsal epitrochlear fossa: (0) present; (1) absent.120. Medial epicondyle orientation: (0) very retroflexed; (1)

slightly retroflexed; (2) medial.121. Medial trochlear keel: (0) anterior to epicondyle; (1)

projects distal to epicondyle.122. Lateral trochlear keel: (0) weak; (1) moderate; (2) strong.123. Entepicondylar foramen: (0) present; (1) absent.124. Olecranon fossa: (0) shallow; (1) deep, sharp laterally.125. Olecranon fossa articular surface: (0) no extension into

fossa; (1) extends into fossa laterally.


126. Trochlear breadth: (0) <capitular; (1) >capitular.127. Distal humerus anteroposterior thickness: (0) very thin;

(1) intermediate; (2) deep.128. Zona conoidea: (0) shallow, broad; (1) deep, narrow.129. Trochlear proximal border: (0) straight; (1) V-shaped for

coronoid beak.130. Humeral shaft curvature: (0) straight; (1) retroflected; (2)

strongly retroflected.131. Deltoid insertion: (0) on proximal half of shaft; (1) on

distal half.132. Humeral head shape: (0) oblong, extending between

tuberosities; (1) less oblong; (2) spherical with tuberositiesanterior.

133. Tuberosity sizes: (0) lesser tuberosity <60% of greatertuberosity; (1) 60e80%.

134. Bicipital groove: (0) broad; (1) intermediate; (2)narrow.

135. Humeral head height: (0) lower than tuberosities; (1)equal or slightly higher; (2) much higher.

136. Humeral/femoral head size: (0) humeral head� femoral;(1) humeral head >femoral.

137. Ulnar olecranon process size: (% of sigmoid notchlength) (0) very short (<50); (1) short (51e80); (2)long (80e100); (3) very long (>100).

138. Olecranon orientation: (0) proximal; (1) posterior.139. Sigmoid notch shape: (0) long, narrow; (1) low, wide.140. Sigmoid median keel: (0) none; (1) strong keel.141. Ulnar shaft cross section: (0) ML< 66% of AP; (1)

ML> 66% of AP.142. Ulnar shaft bowing: (0) convex dorsally; (1) concave or

straight dorsally.143. Distal ulna shape: (0) ML narrow; (1) ML broad.144. Styloidetriquetral articulation: (0) large styloid and

articular surface; (1) small styloid and articularsurface.

145. Distal radioulnar articulation: (0) proximodistally narrow;(1) proximodistally expanded.

146. Distal ulnar facet on radius: (0) faces medially; (1) facesproximomedially.

147. Radial notch position: (0) anterior to lateral third of shaft;(1) anterior to middle of shaft.

148. Radial notch orientation: (0) anterolateral; (1) lateral.149. Radial head outline: (0) oval with flat posterolateral area;

(1) smaller flat area; (2) round.150. Radial head lateral lip: (0) large; (1) present; (2) absent.151. Radial notch shape: (0) single oval facet; (1) two faces at

90-degree angle; (2) two separated facets.152. Radial styloid process: (0) prominent; (1) reduced.153. Radial lunate articular surface: (0) AP narrow; (1)

ventrally expanded.154. Dorsal ridge on radius: (0) absent; (1) present.155. Distal metacarpal shape: (0) widest palmerly; (1)

quadrilateral.156. Tibial condylar facets: (0) symmetrical; (1) medial larger;

(2) lateral larger.157. Distal fibula: (0) robust, flares laterally; (1) reduced, less

lateral flare.

158. Femoral head shape: (0) spherical and separate fromneck; (1) blends with neck superoposteriorly.

159. Femoral head height: (0) below trochanter; (1) at orabove trochanter.

160. Femoral neck tubercle: (0) present; (1) absent.161. Gluteal tuberosity: (0) absent; (1) ridgelike; (2)

distinct.162. Femoral condyles: (0) nearly symmetrical; (1) medial

larger.163. Patellar groove: (0) narrow, deep; (1) broad, shallow; (2)

wider than shaft.164. Talar proximal tubercles: (0) moderately developed; (1)

poorly developed.165. Cotylar fossa: (0) distinct cup; (1) shallow.166. Talar trochlear wedging: (0) strong; (1) weak.167. Talar trochlear rim asymmetry: (0) subequal height; (1)

lateral keel taller; (2) much taller.168. Talar head: (0) width¼ height; (1) width> height.169. Fibular facet of talus: (0) vertical; (1) laterally

projecting.170. Curve of medial trochlear margin (talus): (0) gently

curved; (1) strongly curved.171. Curve of talar body: (0) tight curve; (1) higher radius of

curvature.172. Ectal facet curve: (0) gently concave; (1) sharply

concave.173. Ectal facet shape: (0) broadest anterolaterally; (1) broad-

est proximomedially.174. Anterior calcaneal length: (0) >one-third total length; (1)

one-third; (2) <one-third.175. Posterior calcaneoastragalar joint: (0) faces medially;

(1) faces dorsally.176. Sustentacular facet: (0) undivided; (1) divided.177. Calcaneocuboid joint: (0) round facet with deep pit; (1)

smaller pit.178. Heel tubercle: (0) present; (1) absent.179. Metatarsal II lateral facet: (0) divided; (1) single.180. EntocuneiformeMT I facet: (0) convex with medial

extension; (1) less convex with slight extension.181. Metacarpal II medial facet: (0) undivided; (1) divided by

ligament pit.182. MC I proximal articulation: (0) no lateral extension; (1)

lateral extension.183. Metacarpal dorsal ridges: (0) absent; (1) present.184. TrapeziumeMC I joint: (0) sellar; (1) modified hinge; (2)

non-sellar.185. Os centrale: (0) free; (1) fused.186. Capitulum: (0) no distal expansion of surface; (1) distal

expansion of articular surface.187. Hamate distolateral edge: (0) uninterrupted; (1) inter-

rupted by ligament pit.188. Scaphoid beak: (0) absent; (1) present.189. Capitate distal surface: (0) concave; (1) biconcave.190. Capitate and hamate shape: (0) long, narrow; (1) proxi-

modistally short.191. Hamate hamulus: (0) not projecting; (1) distally

projecting.


Character matrix

‘‘?’’¼missing data; ‘‘-’’¼ inapplicable

Aotus trivirgatus 1 0 0 0 2 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 1 1 1 0 0 1 1 1 0 0 0 0 0 1 0 3 1 1 2 4 1 0 1 0 0 - 0 0 0 3 0 1 2 1 0 2 0 0 1 0 1 0 0 1 1 2 0 0 - 1 - 1 1 0 0 0 1 1 1 1 1 0 1 1 0 2 0 1 1 0 1 1 1 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 1 0 0 1 1 2 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 0 0 2 0 1 1 0 2 0 1 0 1 1 0 0 0 0 0 0 1 0 0 0 1 1 ? 0 ? 0 0 1 0 0 0 0 0 0 0

Saimiri sciureus 0 0 0 0 2 0 0 0 0 0 0 0 0 - 0 0 0 0 0 0 0 0 1 0 0 1 0 1 0 0 0 0 0 1 0 3 0 1 2 4 0 0 1 0 0 - 0 0 0 3 0 1 3 1 0 3 0 0 0 0 1 0 0 1 2 2 0 0 - 1 - 1 1 0 0 0 0 0 0 2 0 1 1 1 0 0 0 1 1 0 1 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0/1 0 0 1 0 0 0 1 0 0 1 1 1 1 1 0 0 0 0 0 0 0 0 0 1 1 0 1 0 0 0 0 0 2 0 0 0 0 2 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 1 1 0 0 0 0 0 1 0 0 0 0 0 0 0

Cebus apella 1 1 0 0 2 0 0 0 0 0 0 1 0 0 0 1 0 0 1 1 0 0 0 0 0 1 1 1 0 0 0 0 0 1 1 3 1 0 1 4 1 1 1 0 0 - 0 0 0 3 0 0 3 1 0 3 0 0 0 0 1 0 0 1 1 2 0 0 - 1 - 1 0 0 1 1 0 0 0 1 0 0 1 1 0 2 0 0 1 0 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 1 0 1 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 1 1 1 1 1 0 0 0 0 0 0 0 0 0 1 1 0 1 0 0 0 0 0 1 0 0 1 0 1 0 2 0 1 1 0 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0

Aegyptopithecus zeuxis 0 0 0 0 2 1 1 0 0 1 1 1 1 - 0 0 1 2 0 1 0 0 0 0 0 0 0 0 1 0 1 0 0 1 0 3 0 0 1 1 0 0 0 0 0 0 1 0 1 3 0 0 2 1 2 3 0 0 0 0 1 2 0 0 0 0 0 1 1 1 0 0 0 0 1 0 0 0 0 0 1 0 1 1 0 0 0 0 0 ? 0 0 0 1 0 0 1 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? 0 0 1 0 0 0 0 0 1 0 0 0 1 0 0 1 0 1 ? 3 0 0 0 ? 0 ? 0 ? ? 0 1 ? ? 0 ? ? 0 ? ? ? 0 1 0 2 0 2 0 0 0 1 1 1 1 1 0 0 0 ? 0 0 ? 1 ? ? ? 0 ? ? 0 ? ? ? ? ?

Dionysopithecus shuangouensis 0 ? ? 1 1 1 1 0 ? ? ? ? ? ? ? ? 1 1 1 1 1 1 1 0 0 0 0 0 2 0 0 0 0 1 0/1 0 0 1 1 2 ? 0 1 0 0 ? 1 0 1 1 0 ? 2 0 1 1 2 1 0 0 1 2 0 0 2 0 0 1 1 1 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

Lomorupithecus harrisoni ? ? ? ? 1 1 1 0 0 1 0 0 0 0 1 2 1 2 1 1 0 ? 0 0 0 0 0 0 1 ? 1 0 0 ? ? 3 ? 0 1 ? ? ? ? ? ? ? ? ? ? ? ? ? 2 0 1 1 0 1 0 0 1 4 0 0 1 0 0 1 2 ? ? ? ? ? ? 1 1 2 1 0 1 1 1 1 ? 0 0 1 ? ? 0 1 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

Epipliopithecus vindobonensis 1 0 0 1 1 1 1 0 0 0 1 0 1 0 0 2 1 2 1 0 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 1 1 2 0 0 0 0 0 1 1 0 1 2 0 0 2 0 0 1 1 0 0 0 1 4 0 0 2 0 0 1 2 0 0 1 1 0 1 1 0 2 1 0 1 1 ? ? ? 0 0 1 1 ? 0 1 0 0 1 1 ? 1 0 0 0 0 0 0 0 0 1 0 ? ? 1 ? ? 0 0 ? 0 0 0 1 0 0 0 0 0 0 1 0 0 0 0 1 1 0 1 ? 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 ? 0 1 0 0 1 2 1 1 1 0 1 1 0 1 0 0 0 1 1 1 0 0 0 0 0 0 1 ? 0 0 ? ? ? ?

Victoriapithecus macinnesi 0 0 0 0 1 1 1 0 0 0 3 1 0 0 2 2 2 0 2 1 2 3 1 0 1 1 1 1 0 0 1 0 0 2 0 2 1 - 1 2 1 1 1 0 0 0 3 0 0 1 0 0 2 1 - 2 0 0 1 1 1 2 - 0 - 2 0 1 1 1 0 2 0 0 ? 1 0 0 0 0 0 0 1 ? 1 2 0 0 0 ? 1 0 1 1 0 0 ? 2 ? ? ? ? ? ? ? ? ? ? 0 2 ? ? ? ? ? 0 ? 0 0 0 1 0 1 1 1 0 2 0 0 2 0 0 1 0 0 ? 3 1 0 0 0 ? 0 0 0 ? 1 0 0 ? 2 ? ? ? 0 ? 1 1 0 ? 0 ? 0 1 ? ? 1 0 0 0 ? 1 ? 0 ? 1 ? 0 ? 1 ? 0 0 0 ? 0 ? 1 ? ? ?

Macaca sp. 0 1 0 0 0 1 1 0 0 0 3 0 0 1 2 2 3 2 2 2 3 3 1 0 1 1 0 1 0 1 1 0 1 2 2 0 0 - 0 ? 1 1 0 1 0 1 3 0 1 2 0 0 0 1 - 1 0 0 1 1 1 - - 1 - 2 0 1 1 1 - 2 0 0 1 1 0 0 0 0 0 0 1 2 1 2 0 0 1 1 1 0 1 1 1 1 1 2 1 0 0 0 0 0 0 0 0 ? 0 2 0 0 0 0 0 0 0 0 1 0 1 0 1 1 1 1 2 0 0 2 0 0 1 1 0 0 3 0 0 0 0 0 0 0 0 0 1 0 0 0 2 0 0 0 0 0 1 1 0 0 0 0 0 1 1 0 2 0 0 0 0 1 1 0 1 1 1 1 0 1 0 0 0 0 0 0 0 1 0 0 0

Proconsul major 0 0 ? 0 1 1 1 0 0 0 1 0 0 - 0 2 1 2 1 0 1 1 0 0 0 0 0 0 2 1 1 0 0 0 0 1 0 0 1 1 1 1 0 0 0 0 2 0 1 3 1 0 2 1 2 2 0 0 1 0 1 2 0 0 0 0 0 0 0 1 1 0 1 ? 0 ? ? ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?Proconsul africanus 0 0 ? 0 1 1 0 0 0 2 1 0 1 1 0 2 1 2 2 0 1 1 0 0 0 0 0 0 2 0 0 0 0 0 0 1 0 0 1 1 1 1 0 0 ? 0 2 0 1 3 1 0 2 1 2 2 0 0 1 0 1 2 1 0 0 1 0 0 0 2 0 0 1 ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

Nyanzapithecus harrisoni ? ? ? ? ? 1 ? ? ? ? ? ? ? ? ? ? ? ? ? 3 3 3 0 1 1 2 0 1 3 0 0 1 0 0 0 1 0 2 2 1 ? ? ? ? ? ? 2 0 1 0 0 1 0 1 2 0 0 0 1 0 1 3 2 0 0 0 1 1 1 1 1 1 ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?


Equatorius africanus 0 0 0 0 1 1 2 1 1 0 1 1 0 1 2 2 1 1 1/2 1 2 3 0 0 0 0 1 1 1 1 1 0 0 1 0 3 0 0 1 1 0 0 1 0 0 0 1 1 1 3 0 0 1 1 2 2 0 0 1 0 1 3 1 0 0 2 0 0 1 1 0 0 0 ? 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? ? ? ? ? ? ? ? ? ? ? 0 1 ? ? ? 1 0 ? 1 0 ? 1 1 1 1 1 ? 1 1 0 1 1 0 0 1 1 1 1 ? 1 1 0 0 0 0 0 1 ? ? 0 ? 1 1 0 ? 0 0 1 0 0 ? 2 0 0 0 1 1 1 ? ? ? ? ? ? ? ? 1 ? 1 1 ? 1 0 0 ? 1 1 ? 1 ?

Afropithecus turkanensis 1 0 0 0 2 1 1 1 1 0 0 0 0 0 ? 0 0 ? 0 1 1 2 0 0 0 1 1 ? 1 1 1 1 0 ? ? 1 ? 1 1 0 0 0 1 0 0 0 1 0 1 3 0 0 ? 1 2 2 0 0 1 0 1 2 1 0 0 1 0 0 1 1 0 0 0 ? 0 1 0 0 0 1 1 0 0 0 1 1 0 0 0 ? 0 0 0 1 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 1 ? ? ? 1 ? ? ? ? ? ? ? ? ? ? ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? 0 ? ? ? ? ? ? 0 0 0 1 ? 1 1 1 ? 0 ? ? ? 0 ? ? ? 1 1 0 0 0 ? ? ? 0 0 ?

Nyanzapithecus vancouveringorum ? ? ? ? ? 1 2 2 2 0 2 1 1 ? ? 0 3 2 0 2 3 3 0 0 0 1 1 1 2 1 0 1 0 1 0 1 0 1 2 0 ? ? ? ? ? ? ? ? ? 3 0 1 0 1 2 0 0 0 1 0 1 3 2 0 0 1 1 1 1 1 ? 0 1 ? 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? 0 1 1 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

Simiolus enjiessi ? ? ? 0 0 1 0 0 0 2 1 0 0 - 1 1 1 0 1 0 1 3 0 0 0 0 0 1 2 1 1 1 0 1 1 3 0 1 2 0 0 0 0 0 0 0 3 0 1 1 0 0 0 1 2 0 0 0 1 0 1 2 2 0 0 1 1 1 1 1 0 0 0 ? 0 1 0 0 0 ? ? ? ? ? ? ? ? 1 ? ? ? ? ? 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 0 0 1 0 0 1 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 1 ? ? ? ? 0 0 1 1 0 ? 1 0 ? ? ? ? ? ? ? ? ? ? 0 ? ? 0 ? ? ? ? ?

Micropithecus clarki 0 0 0 0 2 1 0 0 0 2 1 0 0 0 0 2 1 2 1 0 2 2 0 0 0 0 0 0 1 0 1 1 0 1 0/1 3 0 1 1 3 0 0 1 0 1 1 3 0 1 2 0 0 1 1 2 2 0 0 0 0 1 2 2 0 0 1 0 0 1 2 0 0 0 ? 1 1 1 1 1 1 ? ? ? ? 1 1 ? 1 ? ? ? 1 ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

Kalepithecus songhorensis 0 0 0 0 1 1 0 0 0 0 2 0 0 1 1 2 2 0 1 1 1 1 0 0 0 0 0 0 2 ? 1 0 0 0 0 3 0 0 1 ? 0 0 1 0 0 0 1 0 1 2 1 0 2 1 2 3 0 0 0 0 1 3 1 0 0 0 0 1 0 1 0/1 0 0/1 ? 0 1 0 ? ? 1 ? ? ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 0 0 0 1 1 1 1 0 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 ?

Limnopithecus legetet 0 0 0 0 1 1 1 0 0 0 1 0 1 0 0 2 1 1 1 0 1 2 0 0 0 0 0 0 2 0 0 0 0 1 1 3 0 0 1 1 1 0 1 0 0 0 1 0 0/1 2 0 1 1 1 2 2 0 0 1 0 1 3 0 0 0 0 0 0 0 1 0 1 1 ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 0 ? ? 0 1 ? 0 0 0 ? ? ? 0 1 ? 0 ? ? ? 0 1 1 1 1 0 1 0 ? 1 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ?

Limnopithecus evansi 0 ? ? 0 1 1 1 0 0 0 1 1 1 - 1 1 1 1 1 1 0 1 0 0 0 1 0 0 2 0 0 0 0 1 0 3 0 0 1 1 0 1 1 0 0 0 2 0 1 1 0 0 1 1 2 2 0 0 1 0 1 2 1 0 0 0 0 1 0 1 0 0 0 ? 0 1 0 ? 1 0 ? ? ? ? 0 ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 ? ? ? ? ? ? ? 0 0 ? ? ? ? ? ? ? 0 1 0 1 ? ? ? 0 1 1 ? 1 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

Dendropithecus macinnesi 0 0 0 0 0 1 0 0 0 1 1 0 0 1 1 2 1 0 1 0 0 1 0 0 0 0 0 0 2 0 0 0 0 1 1 0 0 0 1 1 1 1 1 0 0 0 3 0 1 2 1 0 1 1 2 1 0 0 1 0 1 3 0 0 0 0 0 1 0 1 0 0 0 ? 0 ? 1 ? ? 0 ? ? ? ? 0 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? ? ? ? ? ? ? ? 0 1 0 0 1 0 0 1 0 0 0 0 0 ? ? 0 ? ? 3 0 1 0 1 ? ? ? ? ? 0 0 0 1 0 ? 0 0 ? ? ? 0 ? ? ? ? ? 1 0 0 1 1 1 1 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? 0 ? ? ? 0 ?

Rangwapithecus gordoni 0 1 ? 0 0 1 1 0 1 0 1 1 1 - 0 0 2 2 0 0 3 3 1 0 0 1 0 0 2 1 ? 1 0 0 0 1 0 1 2 0 0 0 1 0 0 1 2 0 1 1 0 0 0 1 2 0 0 0 1 0 1 3 0 0 0 0 1 0 0 1 0 0 1 ? 0 ? 1 ? ? 1 ? ? ? ? ? 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 ? ? ? ? ? ? ? 0 1 1 0 ? ? ? 0 0 1 1 1 0 1 0 ? ? 1 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ?

Turkanapithecus kalakolensis ? ? ? ? 1 1 1 2 1 0 2 1 0 - 0 1 2 2 0 ? 2 3 1 1 0 1 0 0 3 1 0 1 0 1 0 2 0 2 1 1 ? ? ? ? ? ? ? ? 1 ? ? ? ? 1 ? 0 0 0 1 0 1 3 1 0 0 1 1 1 1 1 ? ? 1 0 1 ? ? 0 0 0 1 0 0 0 1 ? ? 0 1 ? 0 0 0 1 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 1 0 1 0 1 1 ? ? 1 0 1 0 1 1 0 0 1 0 ? ? ? 0 1 1 1 1 1 0 0 ? 1 1 1 1 0 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

Nyanzapithecus pickfordi 1 1 0 0 0 1 2 2 2 0 3 1 1 1 0 2 3 2 0 3 3 3 1 1 1 2 1 1 3 1 0 1 0 1 0 2 1 2 2 0 1 1 1 0 0 ? ? ? 1 0 0 1 0 1 2 0 0 0 ? 0 1 2 ? 0 0 1 1 1 1 1 1 1 ? ? ? ? 0 ? ? 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?


Gorilla gorilla 1 1 1 0 0 1 1 0 0 0 2 0 0 1 2 1 2 1 1 0 2 3 1 0 1 1 1 1 1 1 0 1 0 0 0 3 0 1 2 3 0 1 1 0 0 0 1 1 1 3 0 1 1 1 2 2 0 0 1 0 1 3 0 0 0 0 1 0 0 1 0 0 0 1 1 1 1 0 0 2 0 0 0 0 1 1 1 0 1 0 0 0 0 1 2 1 2 2 2 1 1 2 1 1 1 1 3 0/2 1 0 1 1 1 1 1 1 1 1 1 1 0 2 1 1 1 1 1 1 1 0 1 2 0 2 1 1 0 0 1 1 1 0 1 1 1 0 0 1 1 2 0 1 1 1 1 1 0 0 1 1 0 1 2 0 0 0 0 1 1 1 1 0 0 2 1 0 1 0 1 0 1 1 1 0 1 1 1 0 1 1 1

Pan troglodytes 1 1 1 0 1 1 1 0 1 0 2 0 0 1 2 2 2 1 2 0 2 2 1 0 1 1 1 1 1 1 0 1 0 1 0 3 0 1 2 2 1 1 1 0 0 0 1 1 1 3 0 0 2 1 2 2 0 0 1 0 1 3 0 0 0 2 0 0 0 1 1 0 0 1 1 1 0 0 0 2 0 1 0 0 1 1 1 0 1 0 0 0 0 1 2 1 2 2 2 1 1 2 1 1 1 1 3 0/2 1 0 1 1 1 1 1 1 0 1 1 2 0 2 1 1 1 1 1 1 1 0 1 2 0 2 1 1 0 1 1 1 1 0 1 1 1 1 0 1 1 2 0 1 1 1 1 1 0 0 1 1 0 1 2 0 0 0 1 1 1 1 1 0 0 2 1 0 0 0 0 0 1 1 1 0 1 1 1 0 1 1 1

Pongo pygmaeus 1 0 1 0 1 1 2 1 0 0 3 1 0 1 2 2 2 1 2 1 2 3 1 0 0 0 2 1 0 1 0 0/1 0 0 0 1 0 1 1 2 1 1 1 0 0 0 1 1 1 3 1 0 2 1 2 3 0 0 1 0 1 3 0 0 0 1 0 0 0 1 0 0 0 1 1 1 0 0 0 2 0 1 0 0 1 1 1 0 1 0 1 0 1 1 0 0 2 2 2 1 1 2 1 1 1 1 3 2 1 0 1 1 1 1 1 1 1 1 1 2 0 2 1 1 1 1 1 1 1 0 1 2 0 2 1 1 0 0/1 1 1 1 0 1 1 1 1 0 1 2 2 0 1 1 0 1 1 0 0 1 1 0 1 2 0 0 0 1 1 1 1 1 0 0 2 0 0 0 0 0 0 1 1 0 0 0 1 1 0 1 1 0

Hylobates sp. 1 1 0 0 0 1 0 0 0 0 3 0 0 1 2 2 3 0 1 0 2 3 1 0 0 0 1 1 1 1 0 0 0 0 0 3 0 1 1 3 1 1 1 0 0 1 2 0 1 1 0 1 1 1 2 2 0 0 1 0 1 3 0 0 0 2 1 0 1 1 1 0 0 1 1 1 1 2 1 2 1 1 0 0 1 0 0 1 1 0 1 1 0 0 1 1 2 2 2 1 0 1 0/1 1 1 1 2 1/2 1 0 1 1 1 1 1 1 0 1 1 2 0 2 1 1 1 1 1 1 1 0 0 2 0 2 1 1 0 0 1 1 1 0 1 1 1 0 0 1 2 2 0 0 1 0 1 0 0 0 1 1 0 0 2 1 0 1 0 0 1 1 1 0 0 1 0 0 0 1 ? 0 1 - 0 2 0 1 ? 0 1 1 0

Presbytis sp. 0 0 0 0 0 1 0 0 0 0 3 0 0 1 2 2 2 0 2 0 3 3 1 0 1 1 0 1 0 1 1 0 1 2 2 0 0 - 0 2 1 1 0 0 0 0 3 0 0/1 2 0 1 1 1 - 2 0 0 1 1 1 - - 1 - 2 1 0 0 1 - 2 0 1 1 1 0 0 1 0 0 1 1 2 1 2 0 1 1 0 1 1 0 1 1 1 2 2 1 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 1 0 1 1 1 0 2 0 0 1 0 ? 1 0 0 0 1 0 0 0 0 1 0 0 0 0 1 0 0 0 1 1 0 0 0 1 1 1 0 1 0 0 0 0 0 1 2 0 0 0 0 0 1 2 ? 1 1 1 0 1 1 0 0 0 0 0 0 1 0 0 0

Cercopithecus sp. 1 0 0 0 0 1 1 2 0 0 3 0 0 1 0/2 2 3 1 2 0 3 3 1 0 1 1 0 1 0 0 0 0 1 2 2 0 0 - 0 3 1 1 0 1 0 1 3 0 1 0 0 0 0 1 - 1 0 0 1 1 1 - - 1 - 2 0 1 1 1 - 2 0 0 1 1 0 0 0 0 0 0 1 2 1 2 0 0 1 1 1 1 1 1 1 1 2 2 1 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 1 0 1 1 1 0 2 0 0 1 0 0 0 1 0 0 3 1 0 0 1 1 0 0 0 0 1 0 0 1 2 0 0 0 0 0 1 1 0 0 0 0 0 1 0 0 2 0 0 0 0 1 1 1 ? 1 1 1 0 1 0 0 0 0 0 0 0 1 0 0 0

Morotopithecus bishopi 1 0 0 ? 2 1 1 1 0 0 1 0 1 - 2 1 0 1 1 1 2 3 1 0 0 0 0 0 2 0 1 0 0 0 1 1 ? 0 1 1 ? ? ? ? ? 0 ? ? 1 ? ? ? 2 1 2 ? 0 0 1 0 1 2 0 0 0 1 0 1 1 ? ? ? ? ? ? 1 0 0 0 1 0 0 0 0 1 1 1 0 0 ? 0 0 0 1 ? ? ? ? 2 1 1 2 1 1 1 1 ? ? ? ? ? ? ? 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 0 ? ? 2 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

Laccopithecus robustus 1 1 0 1 2 1 1 1 1 0 1 0 1 0 0 2 1 2 1 0 1 1 0 0 0 0 1 0 1 0 0 0 0 1 0 0 0 0 1 2 0 0 1 0 0 1 1 0 0 1 0 0 2 0 0 1 0 0 0 0 1 3 1 0 2 1 0 1 2 0 0 0 ? ? ? 1 0 2 1 0 ? 1 ? 1 ? ? 0 1 1 ? ? 1 0 0 1 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

Catopithecus browni 0 1 0 0 2 1 0 0 0 2 3 - 1 - 0 1 1 0 1 0 0 0 1 1 0 0 0 0 1 0 0 1 0 1 - 3 - 0 1 3 0 0 ? 0 0 1 1 0 1 0 0 1 1 0 0 1 0 0 0 0 0 1 1 0 2 2 1 0 1 0 1 0 1 0 0 0 0 0 1 1 1 0 1 1 ? 0 0 0 ? ? ? 1 0 0 0 0 1 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 0 0 0 0 0 1 0 0 ? 1 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 1 1 2 ? ? 0 0 0 1 1 1 1 1 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

Pliopithecus antiquus ? ? ? 1 ? 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 1 0 0 0 1 0 1 ? 0 0 2 0 0 2 2 1 0 0 1 4 2 0 2 0 0 1 2 0 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?

Proconsul nyanzae 0 0 0 0 1 1 0 0 0 0 1 0 0 1 1 2 1 0 1 1 1 2 0 0 0 0 0 0 2 1 1 1 0 0 1 3 0 0 1 1 0 0 ? 0 0 0 2 0 1 3 0 0 2 1 2 2 0 0 1 0 1 3 0 0 0 0 0 0 0 1 0 0 1 ? 1 1 0 ? 0 0 1 0 0 0 ? 1 0 0 ? ? ? 0 0 1 ? ? ? ? 0 0 0 0 0 0 0 0 ? ? 0 ? 1 ? ? ? ? ? ? ? ? ? ? ? 1 1 1 ? 1 1 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 1 0 1 0 1 0 0 0 1/2 1 1 0 0 1 0 ? 0 0 0 1 ? 0 ? 1 ? ? 0 1 0 ? 0 ? 0

Proconsul heseloni 0 0 0 0 1 1 1 0 0 0 1 0 0 1 1 2 1 0 2 1 1 1 0 0 0 0 0 1 1 0 0 0 0 0 0 3 0 0 1 1 0 0 ? 0 0 0 2 0 1 3 1 0 2 1 2 1 0 0 1 0 1 3 0 0 0 0 0 0 1 1 0 0 0 0 1 1 0 0 0 0 1 0 0 0 1 0 0 0 1 ? 0 0 0 1 1 1 2 2 0 0 0 0 0 0 0 0 ? ? ? ? 1 0 0 0 0 0 1 0 1 1 0 1 1 1 1 1 1 1 0 1 0 1 ? 0 ? ? ? 1 0 0 1 1 1 0 0 0 0 1 1 1 0 0 1 0 0 ? 0 ? ? ? 0 0 1 0 0 0 1 1 1 0 0 0 1 0 1 0 0 1 0 0 0 1 0 0 0 1 0 1 0 0 0


References

Andrews, P., 1978. A revision of the Miocene Hominoidea of East Africa.

Bull. Br. Mus. Nat. Hist., Geol. Ser. 30, 85e224.

Andrews, P.J., Harrison, T., Delson, E., Bernor, R.L., Martin, L., 1996. Distri-

bution and biochronology of European and Southwest Asian Miocene cat-

arrhines. In: Bernor, R.L., Fahlbusch, V., Mittmann, H. (Eds.), The

Evolution of Western Eurasian Neogene Mammalian Faunas. Columbia

University Press, New York, pp. 168e207.

Barrett, M., Donoghue, M.J., Sober, E., 1991. Against consensus. Syst. Zool.

40, 486e493.

Begun, D.R., 1989. A large pliopithecine molar from Germany and some notes

on the Pliopithecinae. Folia Primatol. 52, 156e166.

Begun, D.R., 2002. The Pliopithecoidea. In: Hartwig, W.C. (Ed.), The Primate

Fossil Record. Cambridge University Press, Cambridge, pp. 221e240.

Benefit, B.R., McCrossin, M.L., 1991. Ancestral facial morphology of Old

World higher primates. Proc. Natl. Acad. Sci. U.S.A. 88, 5267e5271.

Benefit, B.R., McCrossin, M.L., 1993. Facial anatomy of Victoriapithecus and

its relevance to the ancestral cranial morphology of Old World monkeys

and apes. Am. J. Phys. Anthropol. 92, 329e370.

Benefit, B.R., McCrossin, M.L., 1997. Earliest known Old World monkey

skull. Nature 388, 368e371.

Bishop, W.W., 1964. More fossil primates and other Miocene mammals from

north-east Uganda. Nature 203, 1327e1331.

Bishop, W.W., Miller, J.A., Fitch, F.J., 1969. New potassium-argon age deter-

minations relevant to the Miocene fossil mammal sequence in East Africa.

Am. J. Sci. 267, 669e699.

Conroy, G.C., 1987. Problems of body-weight estimation in fossil primates.

Int. J. Primatol. 8, 115e137.

Delson, E., Andrews, P.J., 1975. Evolution and interrelationships of the catar-

rhine primates. In: Luckett, W., Szalay, F. (Eds.), Phylogeny of the Primates:

A Multidisciplinary Approach. Plenum Press, New York, pp. 405e446.

Fleagle, J.G., 1975. A small gibbon-like hominoid from the Miocene of

Uganda. Folia Primatol. 24, 1e15.

Fleagle, J.G., Simons, E.L., 1978. Micropithecus clarki, a small ape from the

Miocene of Uganda. Am. J. Phys. Anthropol. 49, 427e440.

Gift, N., Stevens, P.F., 1997. Vagaries in the delimitation of character states in

quantitative variationdan experimental study. Syst. Biol. 46, 112e125.

Harrison, T., 1982. Small-bodied apes from the Miocene of East Africa. Ph.D.

Dissertation, University College London.

Harrison, T., 1987. The phylogenetic relationships of the early catarrhine

primates: a review of the current evidence. J. Hum. Evol. 16, 41e80.

Harrison, T., 1988. A taxonomic revision of the small catarrhine primates from

the early Miocene of East Africa. Folia Primatol. 50, 59e108.

Harrison, T., 2002. Late Oligocene to middle Miocene catarrhines from Afro-

Arabia. In: Hartwig, W.C. (Ed.), The Primate Fossil Record. Cambridge

University Press, Cambridge, pp. 311e338.

Harrison, T., Gu, Y., 1999. Taxonomy and phylogenetic relationships of early

Miocene catarrhines from Sihong, China. J. Hum. Evol. 37, 225e277.

Harrison, T., Delson, E., Jian, G., 1991. A new species of Pliopithecus from the

middle Miocene of China and its implications for early catarrhine zooge-

ography. J. Hum. Evol. 21, 329e361.

Kay, R.F., Fleagle, J.G., Simons, E.L., 1981. A revision of the Oligocene apes

of the Fayum Province, Egypt. Am. J. Phys. Anthropol. 55, 293e322.

Kelley, J., 1986. Species recognition and sexual dimorphism in Proconsul and

Rangwapithecus. J. Hum. Evol. 15, 461e495.

Kordos, L., Begun, D.R., 2000. Four catarrhine crania from Rudabanya. Am. J.

Phys. Anthropol. 30 (Suppl.), 199.

Leakey, R.E., Leakey, M.G., 1987. A new Miocene small-bodied ape from

Kenya. J. Hum. Evol. 16, 369e387.

Ness, G., Levi, S., Crouch, R., 1980. Marine magnetic anomaly time scales

for Cenozoic and late Cretaceous: a precis, critique, and synthesis. Rev.

Geophys. Space Phys. 18, 753e770.

Pan, Y., 1988. Small fossil primates from Lufeng, a latest Miocene site in

Yunnan Province, China. J. Hum. Evol. 17, 359e366.

Pickford, M., Senut, B., Gommery, D., Musiime, E., 2003. New catarrhine

fossils from Moroto II, early middle Miocene (ca 17.5 Ma) Uganda. C. R.

Paleovol. 2, 649e662.

Pickford, M.H., 1981. Preliminary Miocene mammalian biostratigraphy for

western Kenya. J. Hum. Evol. 10, 73e98.

Pilbeam, D.R., Walker, A.C., 1968. Fossil monkeys from the Miocene of

Napak, north-east Uganda. Nature 220, 657e660.

Pogue, M.G., Mickevich, M., 1990. Character definitions and character

state delineation: the bete noire of phylogenetic inference. Cladistics 6,

319e361.

Rae, T.C., 1999. Mosaic evolution in the origin of the Hominoidea. Folia

Primatol. 70, 125e135.

Rose, M.D., Leakey, M.G., Leakey, R.E.F., Walker, A., 1992. Postcranial

specimens of Simiolus enjiessi and other primitive catarrhines from the

early Miocene of Lake Turkana, Kenya. J. Hum. Evol. 22, 171e237.

Rossie, J.B., 2005. Anatomy of the nasal cavity and paranasal sinuses in

Aegyptopithecus and early Miocene African catarrhines. Am. J. Phys.

Anthropol. 126, 250e267.

Senut, B., Pickford, M.H., Gommery, D., Kunimatsu, Y., 2000. Un nouveau

genre d’homino€ıde du Miocenene inferieur d’Afrique orientale: Ugandapi-

thecus major (Le Gros Clark & Leakey, 1950). C. R. Acad. Sci., Paris 331,

227e233.

Simons, E.L., 1987. New faces of Aegyptopithecus from the Oligocene of

Egypt. J. Hum. Evol. 16, 273e289.

Slowinski, J., 1993. ‘‘Unordered’’ versus ‘‘ordered’’ characters. Syst. Biol. 42,

155e165.

Steiper, M.E., Young, N.M., Sukama, T.Y., 2004. Genomic data support

the hominoid slowdown and an early Oligocene estimate for the hominoid-

cercopithecoid divergence. Proc. Natl. Acad. Sci. U.S.A. 101, 17021e17026.

Swofford, D.L., 1993. PAUP: Phylogenetic Analysis Using Parsimony Edition

3.1.1. Illinois Natural History Survey, Champaign.

Vogel, C., 1968. The phylogenetical evolution of some characters and some

morphological trends in the evaluation of the skull in catarrhine primates.

In: Chiarelli, A.B. (Ed.), Taxonomy and Phylogeny of Old World Primates.

Rosenberg & Sellier, Turin, pp. 21e55.

Ward, S.C., Kimbel, W.H., 1983. Subnasal alveolar morphology and the sys-

tematic position of Sivapithecus. Am. J. Phys. Anthropol. 61, 157e171.

Ward, S.C., Pilbeam, D.R., 1983. Maxillofacial morphology of Miocene hom-

inoids from Africa and Indo-Pakistan. In: Ciochon, R., Corruccini, R.

(Eds.), New Interpretations of Ape and Human Ancestry. Plenum Press,

New York, pp. 211e238.

Wiens, J.J., 2001. Character analysis in morphological phylogenetics:

problems and solutions. Syst. Biol. 50, 689e699.

Zapfe, H., 1960. Die Primatenfunde aus der Miozanen Spaltenfullung von

Neudorf an der March (Devinzka Nova Ves), Tchechoslowakei. Schwei-

zerische Paleontologische Abhandlungen 78, 4e293.

A new pliopithecoid genus from the early Miocene of Uganda

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