Didier Merle

Emmanuel Côtez

Christine Argot

Henning Blom

Jean Broutin

Gaël Clément

Gregory D Edgecombe

Jin Meng

Brigitte Meyer-Berthaud

Isabelle Rouget

Dominique Gommery
email: dominique.gommery@sorbonne-universite.fr

Lazarus Kgasi
email: lkgasi@ditsong.org.za

Nonhlanhla Vilakazi
email: nvilakazi@uj.ac.za

Frank Sénégas
email: frank.senegas@sorbonne-universite.fr

Martin Pickford
email: martin.pickford@mnhn.fr

Brian Kuhn
email: kuhnbf@gmail.com

Johann Schnyder
email: johann.schnyder@sorbonne-universite.fr

Clés Mots

Pliocène

Véronique Pois
email: veronique.pois@univ-perp.fr

Pr Antiquus T Tp9

T 17

T 20

P Izodi

T T 10

T 13

T 18

P Angusticeps Ka

Keywords: Pliocene, Primates, Parapapio, Cercopithecoides, Cradle of Humankind, Bolt's Farm Primates, Parapapio, Cercopithecoides

 Cercopithecoides cf. williamsi Mollett, 1947. At BPB, we have potentially the oldest known co-occurrence of these two taxa in Southern Africa, the second oldest being at Sterkfontein-Member 2. The currently available evidence suggests dispersal of these two genera from East Africa to Southern Africa, a hypothesis generally accepted by most authors, even though the period of dispersal is now considered to be somewhat earlier than previously thought. Whereas the Early Pliocene environment in the Cradle of Humankind seems to have been drier than that of today, it was probably more humid during the Middle Pliocene which would have allowed the dispersal of these two taxa.

RÉSUMÉ

, et Cercopithecoides cf. williamsi Mollett, 1947. A BPB, nous avons potentiellement la plus ancienne cooccurrence connue de ces deux taxons en Afrique australe, la deuxième plus ancienne étant celle de Sterkfontein-Member 2. Cette découverte suggère une dispersion de ces deux genres de l'Afrique de l'Est vers l'Afrique australe, une hypothèse généralement acceptée par la plupart des auteurs, à une période un peu plus ancienne que l'on pensait jusque-là. Alors que l'environnement du Pliocène inférieur dans le Cradle of Humankind semble avoir été plus sec que celui d'aujourd'hui, il était probablement plus humide au Pliocène moyen, ce qui aurait permis la dispersion de ces deux taxons.

First in situ middle Pliocene cercopithecoid fossils from the Palaeokarst System of Bolt's Farm (South Africa) INTRODUCTION South African karstic palaeontological sites have yielded an abundant collections of cercopithecoid remains [START_REF] Jablonski N | Cenozoic Mammals of Africa[END_REF][START_REF] Bento Da Costa L | Les primates nonhumains pliocènes et plio-pléistocènes d'Afrique du Sud[END_REF] dated to the terminal Pliocene (corresponding to the final late Piacenzian [START_REF] Cohen K | The ICS International Chronostratigraphic Chart[END_REF]) and the early Pleistocene (including here both the Gelasian and the Calabrian [START_REF] Cohen K | The ICS International Chronostratigraphic Chart[END_REF]). This is the case for the "historical sites" of Taung [START_REF]Note on the occurrence of a species of baboon in limestone deposits near Taung[END_REF][START_REF] Gear | A preliminary account of the baboon remains from Taungs[END_REF][START_REF] Freedman L | The fossil Cercopithecoidea of South Africa[END_REF][START_REF] Freedman L | Fossil and subfossil Primates from the limestone deposits at Taung, Bolt's Farm and Witkrans, South Africa[END_REF][START_REF] Jablonski N | Cenozoic Mammals of Africa[END_REF][START_REF] Gilbert | Cercopithecoid humeri from Taung support the distinction of major papionin clades in the South African fossil record[END_REF] which also yielded the first australopithecine (Australopithecus africanus Dart, 1925) and Makapansgat Limeworks (Mollett van der Spuy 1947;[START_REF] Freedman L | The fossil Cercopithecoidea of South Africa[END_REF][START_REF] Maier | New fossil Cercopithecoidea from the lower Pleistocene cave deposits of the Makapansgat Limeworks, South Africa[END_REF][START_REF] Reed K. E | The Paleoecology of Makapansgat and Other African Plio-Pleistocene Hominids Localities[END_REF][START_REF] Jablonski N | Cenozoic Mammals of Africa[END_REF] Most of the fossiliferous deposits in the Sterkfontein Valley are remnants of karstic systems [START_REF] Brain | The Transvaal Ape-Man-Bearing Cave Deposits[END_REF](Brain , 1981) ) mined for speleothems by the gold industry in the 19 th and the beginning of the 20 th century. What the miners regarded as waste, such as blocks of fossiliferous breccia, was discarded onto dumps. The largest fraction of Plio-Pleistocene (classically covering the period 3.1-0.9 Ma in South Africa) fossil material originates from such mine dumps. The second largest fraction originates from decalcified breccia pockets or makondoes [START_REF] Reynolds S | Sterkfontein at 75: review of palaeoenvironments, fauna and archaeology from the hominin site of Sterkfontein (Gauteng Province, South Africa)[END_REF][START_REF] Adams | A multi-disciplinary perspective on the Age of Australopithecus in Southern Africa[END_REF]. A small portion of the fossils are obtained via collecting of specimens from in situ breccia, providing a better stratigraphic context than the dump waste and decalcified pockets. On Bolt's Farm more than thirty fossil-bearing localities have been identified [START_REF] Cooke H | Dinofelis barlowi (Mammalia, Carnivora, Felidae) cranial material from Bolt's Farm, collected by the University of California African Expedition[END_REF][START_REF] Gommery D. & Braga | Palaeontological sites on 'Bolt's Farm', Sterkfontein Valley, South Africa[END_REF][START_REF] Claude | a. -Plio-Pleistocene fossils from the Femur Dump, Bolt's Farm, Cradle of Humankind World Heritage Site[END_REF][START_REF]Preliminary results concerning the discovery of new fossiliferous sites at Bolt's Farm (Cradle of Humankind, South Africa)[END_REF]Gommery et al. , 2016;;[START_REF] Monson T | Materials collected by the southern branch of the UC Africa Expedition with a report on previously unpublished Plio-Pleistocene fossil localities[END_REF][START_REF] Gommery D | Fossil Suidae (Artiodactyla, Mammalia) from Aves Cave I and nearby sites in Bolt's Farm Palaeokarst System, South Africa[END_REF], 2020;[START_REF] Edwards T. R | Combining legacy data with new drone and DGPS mapping to identify the provenance of Plio-Pleistocene fossils from Bolt's Farm, Cradle of Humankind (South Africa)[END_REF], ranging from the early Pliocene to the end of the early Pleistocene, some of which have yielded cercopithecoid fossils [START_REF] Freedman L | The fossil Cercopithecoidea of South Africa[END_REF][START_REF] Freedman L | Some news fossil, Cercopithecoid specimens from Makapansgat, South Africa[END_REF][START_REF] Freedman L | Fossil and subfossil Primates from the limestone deposits at Taung, Bolt's Farm and Witkrans, South Africa[END_REF][START_REF] Freedman L | A new check-list of fossil Cercopithecoidea of South Africa[END_REF][START_REF] Freedman L | South African fossil Cercopithecoidea: A re-assessment including a description of new material from Makapansgat, Sterkfontein and Taung[END_REF][START_REF]Evolutionary History of the Primates[END_REF][START_REF]Cercopithecid biochronology of the African Plio-Pleistocene: correlation among eastern and southern hominid-bearing localities[END_REF]El-Zaatari et al. 2005;Gilbert 2007a, b;Williams et al. 2007;[START_REF] Kuykendall K. L. & Rae T | Presence of the maxillary sinus in fossil colobinae (Cercopithecoides williamsi), from South Africa[END_REF]Gommery et al. 2008a[START_REF]Cercopithecoidea material from the Middle Pliocene site, Waypoint 160, Bolt's Farm, South Africa[END_REF]Gommery et al. , 2016;;[START_REF] Claude | a. -Plio-Pleistocene fossils from the Femur Dump, Bolt's Farm, Cradle of Humankind World Heritage Site[END_REF][START_REF] Jablonski N | Cenozoic Mammals of Africa[END_REF][START_REF] Gilbert | Evolution of the modern baboon (Papio hamadryas): A reassessment of the African Plio-Pleistocene record[END_REF]. The oldest cercopithecoid from Bolt's Farm, dating to the early Pliocene, came from the locality of Waypoint 160 (Gommery et al. 2008b(Gommery et al. , 2009[START_REF]Cercopithecoidea material from the Middle Pliocene site, Waypoint 160, Bolt's Farm, South Africa[END_REF][START_REF] Brink | Waypoint 160, Bolt's Farm Cave System: First in situ primate remains[END_REF]. The locality of Waypoint 160 has been dated biochronologically through the identification of associated rodents, such as Euryotomys (Sénégas [START_REF] Avery | -New evidence for the murine origins of the Otomyinae (Mammalia, Rodentia) and the age of Bolt's Farm (South Africa)[END_REF][START_REF] Michaux | Boltimys broomi gen. nov., sp. nov. (Rodentia, Mammalia), nouveau Muridae d'affinité incertaine du Pliocène inférieur d'Afrique du Sud[END_REF]. Whereas almost all the material of Cercopithecoidea from this locality came from blocks of breccia in the mine dumps, a fragment of a tooth was found in situ [START_REF] Brink | Waypoint 160, Bolt's Farm Cave System: First in situ primate remains[END_REF]. In contrast to East Africa and except for the late Piacenzian, the Pliocene cercopithecoid fossil record in Southern Brad Pit or BP (with its two loci Brad Pit A or BPA and Brad Pit B or BPB) is a fossiliferous site discovered on the 29 th of November 2010 on Klinkert's portion of Bolt's Farm [START_REF]Preliminary results concerning the discovery of new fossiliferous sites at Bolt's Farm (Cradle of Humankind, South Africa)[END_REF]) (Fig. 1). An outcrop of in situ breccia at BPB yielded some specimens of a cercopithecoid associated with a rich mammal microfauna. The preliminary study of the rodents suggests a middle Pliocene correlation. BPB probably represents the second oldest fossiliferous deposit in the Cradle of Humankind as well as the oldest co-occurrence of the two genera Parapapio [START_REF] Jones T. R | A new fossil primate from Sterkfontein, Krugersdorp, Transvaal[END_REF], and Cercopithecoides Mollett, 1947, in South Africa (Jones 1937;Mollett Van Der Spuy 1947). Member 2 of Sterkfontein yielded some remains of these two genera and may be of similar age [START_REF] Pickering T. R | The context of Stw 573, an early hominid skull and skeleton from Sterkfontein Member 2: taphonomy and paleoenvironment[END_REF][START_REF] Granger D. E | New cosmogenic burial ages for Sterkfontein Member 2 Australopithecus and Member 5 Oldowan[END_REF]. This article describes the first craniodental and postcranial remains of the cercopithecoids from BPB, retrieved from in situ breccia. This study increases our knowledge about the Cercopithecoidea of the middle Pliocene in South Africa, which are rare, and will contribute to a better understanding of their evolutionary history.

GEOLOGICAL AND PALAEONTOLOGICAL CONTEXTS

Until recently, the two loci BPA and BPB were considered to represent different karst infills (Fig. 1). The excavations done since 2013 have exposed more geological details since the discovery of the locality. A preliminary geological study done in 2019 suggests that the two loci represent in fact a single infill. BP consists of an un-roofed section of palaeokarst infill hosted in Palaeoproterozoic dolomites of the Malmani Subgroup (Chuniespoort Group, Transvaal Supergroup). The effects of the past mining activity at BPB, as well the western part of BPA, were less intense than at some other loci at Bolt's Farm. The primate material described herein comes from the north-western corner of BPB and was retrieved from the top section of the exposed breccia, informally referred to as the "Parapapio spot" (Fig. 2). At the time of the discovery of BPB on the 29 th of November 2010, a tooth and some fragments of bones were noticed at the top of this spot (Fig. 3) which, after acid preparation are in fact parts of a skull of Parapapio Jones, 1937.

The excavation activities have confirmed the in situ context of "Parapapio spot" as well as two other important outcrops of breccia at BPB. It was important to confirm the in situ context of the "Parapapio spot" because the geology of the palaeokarstic sites in the Sterkfontein Valley is complex and often disrupted by past mining activities. The preliminary geological study (Fig. 4A) recognized three main sedimentary units, ranging between 3.25 and 3.75 m thick. The "Parapapio spot" belongs to Unit 2 (0.85 m thick) (Fig. 4B) which has a finer matrix, when compared to Unit 1, comprised of medium sands with conglomeratic debris. The whole deposit consists of a fining-upward sequence with conglomeratic levels at the base changing to sand/silt at the top (Fig. 4A). Unit 2, like Unit 1, shows numerous fluid escape figures and locally fossiliferous accumulations. They correspond to the sedimentary filling of the karst cavity.

Some breccia samples taken from the top of the "Parapapio spot" were prepared by chemical means (see material and method). Associated with the primate fossils, there are other macrofaunal remains which include a few post-cranial bones of a large felid and of a medium-sized bovid, which are currently under study. A well-preserved cranial fragment of a bald ibis, Geronticus thackerayi Pavia, 2019, was also extracted [START_REF] Pavia | Geronticus thackerayi, sp. nov. (Aves, Threskiornithidae), a new ibis from the hominin-bearing locality of Kromdraai (Cradle of Humankind, Gauteng, South Africa)[END_REF]Pavia et al. 2017). In contrast, the microfaunal component is

MATERIAL AND METHODS

The BPB specimens are housed in the Plio-Pleistocene Palaeontology section at the Ditsong: National Museum of Natural History (DNMNH) in Pretoria, South Africa. With these fossil specimens being very fragile and fragmentary, it was decided that the most controlled method of preparation of the surrounding breccia matrix, was chemical preparation (6% solution of acetic acid and protected with Paraloid B-72 consolidant) [START_REF]The effectiveness of fossil acid preparation at Ditsong: National Museum of Natural History, South Africa[END_REF]. Upper case letters are used for upper teeth and lower case for lower teeth, followed by the meristic position (see abbreviations). To prevent confusion due to the diversity of dental nomenclatures employed in the literature, a "/" is placed after the letter and the meristic position for upper teeth (e.g. M1/, upper first molar) and the "/" is placed between the letter and meristic position for lower teeth (e.g. p/3 lower third premolar) the forward slash representing the occlusal plane. The deciduous stage of the tooth is indicated by a "d" placed before the tooth. The dental anatomical nomenclature is based on and adapted from [START_REF] Gommery | Evolutionary History of the Cercopithecidae[END_REF][START_REF]Primate Dentition. An Introduction to the Teeth of Non-human Primates[END_REF] and [START_REF]Evolutionary History of the Primates[END_REF]. Dimensions of the teeth are given in millimetres and follow the method of [START_REF] Freedman L | The fossil Cercopithecoidea of South Africa[END_REF], in particular for the mesial (Mb) and distal (Dd) widths of the molars.

The present study corresponds to anatomical descriptions with some linear measurements in mm taken with a manual plastic calliper with an accuracy of 0. 1). The systematics of the smallest South African fossils of Papio Erxeleben, 1777, change regularly, and in this paper we follow the one adopted by [START_REF] Gilbert | Evolution of the modern baboon (Papio hamadryas): A reassessment of the African Plio-Pleistocene record[END_REF]. The specimens from Taung are considered to be Papio izodi [START_REF] Gear | A preliminary account of the baboon remains from Taungs[END_REF], and the others from Kromdraai A and Cooper's are Papio angusticeps (Broom, 1940). For the linear dental measurements concerning the specimens from BPB attributed to Parapapio broomi and Papionina indet., data from the literature were added to our measurements [START_REF] Freedman L | The fossil Cercopithecoidea of South Africa[END_REF][START_REF] Freedman L | Fossil and subfossil Primates from the limestone deposits at Taung, Bolt's Farm and Witkrans, South Africa[END_REF][START_REF] Frost S. R | -New Early Pliocene Cercopithecidae (Mammalia: Primates) from Aramis, Middle Awash Valley, Ethiopia[END_REF][START_REF] Frost S. R. & Delson E | -Fossil Cercopithecidae from the Hadar Formation and surrounding areas of the Afar Depression, Ethiopia[END_REF] to increase the sample (Table 2; Appendices 1-7 3,5). -Mandibles. BPB 2 (Fig. 11; Tables 5,6); BPB 3 (Fig. 14; Tables 5,6); BPB 4 (Fig. 9); BPB 13 (Fig. 16; Table 6); BPB 30 (Fig. 17; Table 6).

desCription Cranium BPB 1 (Figs 5,6; Tables 3,5) General description and preservation. This fossil is a highly distorted and crushed partial skull (right side of the skull and part of the left side of the face) (Fig. 5A-C). It lacks an important part of its left side probably due to erosion of the "Parapapio spot". At the time of discovery, the skull was partially exposed and the right side of the skull was protected in the breccia (Fig. 4). Due to the fossilisation, the skull is almost flattened, the right part of the calvarium is practically in the same plane as the face, which complicates the anatomical study according to the classical anatomical plans. The fossil is 110.4 mm long antero-posteriorly (from the broken anterior part of the muzzle to the preserved posterior part of the skull). The face is 92.8 mm long (from the broken anterior part of the muzzle to the glabella) and the preserved width is approximately 53.6 mm (from the broken lateral edge of the left orbit to the lateral edge of the right orbit). The preserved part of the maxilla is approximately 50 mm long anteroposteriorly (from the fragment of the P3/ to the end of the bone of the dental arch posteriorly to the M3/). The classical anatomical landmarks are not preserved here for the linear measurements and it is therefore impossible to make a further comparison with data from the literature or on more complete specimens. Much of the back of the skull is not preserved and originally the skull was longer, the muzzle is relatively short if we compare it with skulls of extinct and extant specimens of Papio, except for Papio izodi (see [START_REF] Gilbert | Evolution of the modern baboon (Papio hamadryas): A reassessment of the African Plio-Pleistocene record[END_REF]. This skull belongs to a medium-sized to large papionine such as Parapapio. The skull also presents several other diagnostic features of Parapapio [START_REF] Freedman L | The fossil Cercopithecoidea of South Africa[END_REF][START_REF] Freedman L | Fossil and subfossil Primates from the limestone deposits at Taung, Bolt's Farm and Witkrans, South Africa[END_REF][START_REF]Evolutionary History of the Primates[END_REF][START_REF] Jablonski N | Cenozoic Mammals of Africa[END_REF][START_REF] Harrison T | Cercopithecids (Cercopithecidae, Primates)[END_REF][START_REF] Gilbert | Cladistic analysis of extant and fossil African papionins using craniodental data[END_REF]) and differs from the smallest taxa of fossil Papio such as P. izodi and P. angusticeps [START_REF] Gilbert | -Papio cranium from the hominin-bearing site of Malapa: Implications for the evolution of the modern baboon cranial morphology and South African Plio-Pleistocene biochronology[END_REF][START_REF] Gilbert | Evolution of the modern baboon (Papio hamadryas): A reassessment of the African Plio-Pleistocene record[END_REF], namely, in lateral view (Fig. 5C), despite some small cracks, the interorbital and nasal regions indicate a slight concavity and lack of an ante-orbital drop. The maxillary ridge is weak in comparison with that of Papio and the maxillary fossa seems shallow. In anterior view (Fig. 5B), as well as in superior view (Fig. 5D), despite the lack of some superficial bone, the supraorbital torus is lightly built and not projected anteriorly, and the glabella is not prominent. In superior view, the ophryonic groove is absent. The anterior temporal line is weak. Only the right upper toothrow (P3/-M3/) is preserved but the P3/ is badly damaged (Figs 5E; 6A, B).

Calvaria.

It is represented mainly by the frontal and a part of its right side (sphenoid, anterior parts of temporal and parietal) (Fig. 5C,D). The frontal presents many cracks and is distorted. It lacks some fragments of bone. A hole located postero-laterally has a rounded aspect and perhaps could be related to a predation mark. Despite the distortion, the observations in the anterior and right lateral views suggest that the anterior part of the frontal was moderately rounded. Only the right inferior temporal line is preserved. This line is well defined (weak crest), short (seems to stop abruptly after the postorbital constriction). The frontal doesn't exhibit an ophryonic groove, just a very shallow depression on the right side and probably on the opposite side but this part is badly damaged. The morphology of the temporal line and the curved aspect of BPB 1 are similar to BF 43 in which the temporal lines are pinched, as well in other skulls attributed to Pp. broomi [START_REF] Gilbert | Cladistic analysis of extant and fossil African papionins using craniodental data[END_REF]. In antero-lateral and lateral views, the sphenoid is covered anteriorly by some breccia and broken posteriorly at the suture with the temporal. The remnant of temporal presents many cracks and is heavily distorted. In its inferior part, a groove marks the beginning of the zygomatic arch.

Basicranium. Only the anterior part of the right side of the basicranium is preserved (Fig. 5E). At the junction with the palate, the lateral pterygoid plate is well preserved, except for a lack of a central fragment, and as well the deep fossa pterygoidea. Laterally, the fossa mandibularis is preserved but is slightly distorted and flattened. Posteriorly, there is a wellpreserved robust posterior glenoid process which is easier to observe in the lateral view.

Orbital region. The face of BPB 1 is badly damaged and distorted but it is possible to estimate the original morphology (Fig. 5A-C). The face lacks some superficial bone but this doesn't modify greatly the morphology but alters the aspect of the supraorbital torus which seems lightly built and not projected anteriorly. There is no important bulge between the two orbits which means that the glabella is not prominent (Fig. 5C). The inter-orbital pillar (between the superior median corners of each orbit) is slightly broad (11.6 mm) as in BF 43 (11.5 mm) and long. The right orbit is more complete (the median and the superior margins, as well the medio-inferior corner, are preserved) than the left where only the median margin and medio-superior corner remain. The preserved morphology of the right orbit (23.2 mm height) seems to suggest that the orbit is squarish with rounded corners as observed in BF 43. The median part of the orbit is mostly filled with matrix but not near the inter-orbital pillar which allows us to observe inferiorly a developed fossa for a well-preserved lacrimal duct. In the medio-superior corner of the margin of the left orbit, there is a small supra-orbital notch (Incisura supraorbitalis). It is impossible to see this anatomical structure in the right orbit because there is a crack in the same area. The right zygomatic process of the maxilla is damaged, but it is possible to observe its origin (or root).

The malar height is short as in BF 43 (the malar height is the distance between the inferior orbital margin and the inferior border of the zygomatic process of the maxilla as defined in [START_REF] Gilbert | -Papio cranium from the hominin-bearing site of Malapa: Implications for the evolution of the modern baboon cranial morphology and South African Plio-Pleistocene biochronology[END_REF]).

Muzzle and palate

The anterior part of the snout is badly damaged (Fig. 5A,B). The premaxilla is not preserved and neither is the nasal aperture. The nasal is preserved only posteriorly and has many cracks. It is slightly distorted but this doesn't affect the morphology. It is convex transversely and slightly concave antero-posteriorly near the level of the inferior edge of the orbits as in BF 43. There is no ante-orbital drop. The root of the right zygomatic process is located above the middle of the M3/ as in BF 43 and suggests that the muzzle is relatively short. The right maxilla is preserved and some fragments of the left maxilla are preserved along the nasal from the orbit to the premaxilla. The right maxilla is distorted but there is no indication of a strong maxillary ridge and of a deep maxillary fossa.

Only the right side of the palate is preserved (Fig. 5E). The postcanine tooth row is slightly curved and the lingual wall of the dental arch is moderately tall which suggest that the palate was moderately deep. Below and lingually to the M3/, there is an antero-posteriorly oriented depression which corresponds to the groove of the posterior palatine foramen. The bone of the dental arch doesn't end immediately posteriorly to the M3/ but presents the same aspect observed in BF 43, forming a robust bony bulge. This suggests that the hard palate ends behind the M/3.

Dentition.

The post-canine teeth are well preserved except for the P3/ which is just represented by a lingual fragment and the P4/ which has lost a fragment on the buccal edge of occlusal surface (Fig. 6 A; Tables 345). The P4/ and M1/ are worn. The M2/ is less worn (more lingually than buccally). The M3/ is slightly worn at the apex of the protocone and the hypocone. The degree of wear of the preserved post-canine teeth is typical of a mature, but not very old, adult. Despite an important MPI (maxillary premolar index [START_REF] Fleagle | Skeletal and dental morphology supports diphyletic origin of baboons and mandrills[END_REF][START_REF] Frost S. R. & Delson E | -Fossil Cercopithecidae from the Hadar Formation and surrounding areas of the Afar Depression, Ethiopia[END_REF] of 65.11 which correspond to the upper variation of Pp. broomi and perhaps the lower variation of Pr. antiquus [START_REF]Note on the occurrence of a species of baboon in limestone deposits near Taung[END_REF] (Gilbert 2007a), the size of this premolar enters into the variation of Pp. broomi and Pp. whitei Broom, 1940, from Sterkfontein and Makapansgat Limeworks (Tables 3,4; Fig. 7). The molars increase in size from the M1/ to the M3/ (Tables 3;4). The range of variation in molar size overlaps with different taxa of fossil South African papionines (Figs 8; 9) but differs from that of Papio jonesi. The size of the two molars of BPB1 plots within the range of variation of Pp. broomi, especially true for the M2/. The M3/s have a reduced distal loph as in BF 43. [START_REF] Heaton | Taxonomy of the Sterkfontein Fossil Cercopithecinae: the Papionini of Members 2 and 4 (Gauteng, South Africa)[END_REF] found this feature in Pp. broomi. The shape outline of the base of the crown is more trapezoidal than for the M2/.

The well-preserved aspect of the occlusal surface of the M3/ permits observation of the intercuspid distance which is small and the molar reversed W-shaped in outline as described by [START_REF] Mckee | Taxonomic and evolutionary affinities of Papio izodi fossil from Taung and Sterkfontein[END_REF]. The M3/ presents an additional cusp linguodistally (Fig. 6A,B), a developed distoconulus [START_REF]Morphological studies of Macaca fuscata. IV Dentition[END_REF] as in an isolated M3/ from BPA (BPA 230 under study) and Sts 378A but less robust in this tooth than specimen BPB 1 (Freedman 1957). This strongly increases the mesio-distal length of the M3/ of BPB 1 (Fig. 10). In the oblique-lingual view (Fig. 6B), there is an additional strong root beneath the distoconulus for BPB 1 and the same applies to BPA 230. This distoconulus increases the length of the dental row P4/-M3/ or M1/-M3/ (Table 3). It seems that no distoconulus is observed in extant Papio (Swindler 2002).

Mandible BPB 2 (Fig. 11; Tables 567) General description and preservation. BPB 2 is a partial right mandible broken mesially to the p/3 and the symphysis is absent (maximum length: 100.4 mm; breadth: 25.4 mm; height: 34.7 mm). The anterior part is badly damaged, and the two premolars are fragmentary. The corpus mandibularis is poorly preserved with some more or Mandibular corpus and ramus. It is poorly preserved on the anterior portion which presents cracking, and some fragments of bone are missing. The mandibular corpus and the gonial region are broken along the inferior edge. Despite this damage, the mandibular corpus is moderately deep (preserved height between m/1-m/2 is 25.4 mm and between the m/2-m/3 is 24.9 mm). Buccally, there is no prominentia lateralis under the m/3 (Fig. 11A is flat and vertical as in Parapapio. Inferiorly the lingual surface (Fig. 11B) presents an important submandibular fossa. The ascending ramus is almost in the alignment of the molar tooth row. In contrast, in BF 42B (C. williamsi), the ascending ramus is not so laterally located. The extramolar sulcus is narrow medio-laterally as well as mesio-distally. The ramus is low and 31.3 mm long antero-posteriorly. The distal edge of the ascending ramus tilts disto-superiorly. On the lingual surface, the mandibular foramen is low, located at the same level as the alveolar margin, and the groove of this foramen is deep and pronounced. The ascending ramus has lost the coronoid process. The mandibular condyle is well preserved and is more developed medially (maximum length: 8.5 mm; breadth: 13.7 mm).

Dentition. The two premolars are very poorly preserved (Fig. 11C). The mesio-buccal flange on the p/3 (Fig. 11A) is the only well-preserved part and is short as in female individuals of Parapapio. The flange is fairly well developed and not honed by the upper canine which suggests that this specimen represents a female. The three molars have lost some fragments of enamel. The molars increase in size from m/1 to m/3 as is usual in papionines (Tables 567).

The m/1 is the least well preserved and the most deeply worn of the three molars (the preservation doesn't permit inclusion in the bivariate plots MD x BL). The molars possess relatively thick enamel and the less worn cusps are low (Fig. 11D). On the m/3, the prominent and salient hypoconulid is located buccally. This continues buccally by a developed distal marginal ridge and the distal fovea is deep, spacious and short mesio-distally as in many specimens of Pp. broomi (such as Sts 414A and Sts 565) and also but sometimes less pronounced in Pp. whitei (such as Sts 563). The dimensions of the two molars enter into the range of variation in molar size of different species of Parapapio but the m/2 doesn't enter in the range of variation of Pp. jonesi .

Mandible BPB 3 (Fig. 14; Tables 567) General description and preservation. BPB 3 represents a left mandible in two pieces, broken between the p/3 and p/4. The smaller fragment (21.9 mm long antero-posteriorly, 19.6 mm broad and 19.3 mm tall under the p/3) corresponds to the left part of the symphysis. The second fragment from the mesial side of the p/4 to the posterior of the ramus is 82.4 mm long antero-posteriorly and 40.1 mm high. The two fragments fit together well. The upper part of the ramus is absent and the gonial region is damaged.

The bone presents some superficial cracking.

Symphysis.

Only the left part of the symphysis is preserved. The inferior edge is broken. In buccal view (Fig. 14A), the symphysis extends at least as far posteriorly as the m/1 (a fragment is preserved at the inferior edge of the mandibular corpus below the contact p/4-m/1). In lingual view, the planum alveolare seems long and slopes moderately disto-inferiorly (Fig. 14B). This is moderately developed transversely and is moderately deep. The superior transverse torus is smooth. The digastric fossa seems small and shallow. The anterior surface is eroded.

Mandibular corpus. The mandibular corpus is better preserved than that of BPB2. It presents some superficial cracking, especially below the m/3. The inferior edge of the mandibular corpus is well preserved. It is moderately deep (height between m/1-m/2 is 25 mm and m/2-m/3 is 24.7 mm) and robust (mid-height width between m/1m/2 is 11.5 mm and m/2-m/3 is 13 mm). The buccal surface (Fig. 14A) is slightly convex and vertical. Buccally, there is a mental foramen under the p/3 and no prominentia lateralis under the m/3. Distally the lingual surface (Fig. 14B) presents a shallow depression (submandibular fossa). The preserved part of the ascending ramus suggests that it is almost in the alignment of the molar tooth row. The extramolar sulcus is narrow medio-laterally as well as mesio-distally.

Dentition. The canine and incisors are absent (Fig. 14C).

The premolars and molars are better preserved than in BPB 2 despite presenting some cracking (especially in p/4, m/1 and m/2) (Fig. 14D). The molars increase in size from m/1 to m/3 as for BPB 2 (Tables 567). The two premolars and the m/1 are deeply worn, m/2 is moderately worn and the m/3 has the two mesial cuspids worn (protoconid and paraconid) which suggest a mature adult individual. The degree of wear of the three molars corresponds to that observed in BPB 2. Probably, BPB 2 and BPB 3 specimens belong to the same individual. The mesio-buccal flange on the p/3 is fairly developed as is typical of females. The hypoconulid is well developed and located buccally as in BPB 2. The aspect of the distal fovea is very similar to that of BPB 2 as well. Concerning the dimensions of the m/2 and the m/3, they are the same as for BPB 2 (Figs 12-13).

Mandible BPB 4 (Fig. 15) General description and preservation. BPB 4 is the distal part of the right mandible (maximum length: 75.4 mm; height: 49.3 mm). The coronoid process is broken as is the mesial edge of the ascending ramus. The fossil presents some cracking, the most important are on the distal part of the ascending ramus. The mandibular condyle is well preserved but the coronoid process is absent. No tooth is preserved, only the sockets for the roots of the m/3. Mandibular corpus and ramus. Only the portion below the m/3 is present and suggests that the mandibular corpus is vertical, high and moderately large (below the alveolar margin at m/3, height: 28.2 mm; breadth: 12.7 mm). The buccal surface (Fig. 15A) is slightly convex and there is no prominentia lateralis under the m/3. The extramolar sulcus is only preserved buccally. Inferiorly the lingual surface (Fig. 15B) presents a depression which corresponds to the submandibular fossa.

Gommery D. et al.

The gonial region is well preserved and only a small fragment of the inferior edge is broken. This region doesn't expand distally or inferiorly but presents an open convex outline. In the lingual surface, at the junction of the gonial region with the distal edge of the ascending ramus, there are three smooth bulges which correspond to medial pterygoid tubercles. The ascending ramus is low and 36 mm long antero-posteriorly. The distal edge of the ascending ramus tilts disto-superiorly. On the buccal surface (Fig. 15A), the mandibular foramen is present and the surface bone covering the canal of this foramen is lacking, probably due to damage during fossilization. The mandibular condyle is well developed medially (maximum length: 7.1 mm; preserved breadth: 17.4 mm). The general morphology of the ascending ramus of BPB 4 is very similar to that of BPB 2.

Dentition. This fragment is edentulous, but the sockets for the roots of the m/3 are present, well preserved for the distal root and partially preserved for the mesial root (Fig. 15A,C).

Mandible BPB 13 (Fig. 16; Tables 6,7) General description and preservation. BPB 13 is the distal part of a left mandible with the m/3 (maximum length: 43.9 mm; height: 42.6 mm). The fossil presents some cracking. The coronoid process is broken and the mandibular condyle is damaged. The gonial region is preserved as is most of the posterior part of the ascending ramus. The tooth row is only represented by the well-preserved m/3. Mandibular corpus. Just a part of the mandibular corpus is preserved below the m/3. The buccal surface (Fig. 16A) is almost flat and there is no prominentia lateralis under the m/3. The extramolar sulcus is short mesio-distally and narrow bucco-lingually as in BPB 3 (Fig. 16C).

The gonial region is well preserved, except for a few fragments of missing bone. The dorso-inferior outline of this region is convex. The lingual surface of the gonial region (Fig. 16B) is slightly concave. The mandibular foramen is well preserved and the mylohyoid notch is also well preserved. The ascending ramus is better preserved in its posterior part and the posterior edge of the ascending ramus tilts slightly dorsally. The anterior edge of this ramus is broken but the ramus is clearly low and long. The mandibular condyle is broken lingually and abraded (maximum preserved length: 5.9 mm and breadth: 9.9 mm).

Dentition. The m/3 is well preserved and just the apex of the two mesial cusps (protoconid and metaconid) are slightly worn (Fig. 16D). This fossil corresponds probably to a younger adult than BPB 2 and BPB 3. The cuspids are more centrally positioned than in Papio robinsoni (laterally positioned). This molar presents different extra-cusp and cusplet. A small Tuberculum intermedium is present between the metaconid and the entoconid. A low accessory cusp, near the base of the crown is also present between the protoconid and the hypoconid. The distal part of the molar is developed with a Tuberculum sextum. The hypoconulid is important and is located disto-lingually. Buccally, there is a clear distal marginal ridge and the distal fovea is deep. The size of the molar plots within the range of variation of Pp. broomi (Tables 45; Fig. 13).

Mandible BPB 30 (Fig. 17; Tables 6,7) General description and preservation. BPB 30 is part of a left mandible with fragmentary m/2 and worn m/3 (maximum length: 44.9 mm; breadth: between m/2-m/3: 10.9 mm; height: 29.3 mm).

Mandibular corpus.

The mandibular corpus is only preserved below the m/2 and the m/3 and is not preserved over the entire height. It seems narrow inferiorly. On the lingual surface (Fig. 17B), a deep concave depression slopes from mesio-inferiorly to disto-superiorly and corresponds to the submandibular fossa. On the buccal surface (Fig. 17A), there is a smooth concavity probably due to fossilization and it seems that the prominentia lateralis doesn't exist. The mandibular corpus presents a slender aspect unlike Cercopithecoides. A short portion of the extramolar sulcus is preserved. This is reduced and located disto-buccally to the m/3 and not buccally as in Cercopithecoides.

The crown of the m/2 is represented only by a fragment of dentine (Fig. 17C). The m/3 is very worn suggesting that it represents an old adult and lingually it lacks some fragments of enamel at the metaconid and entoconid. The enamel is thick and the median buccal and lingual clefts are narrow as usual in Papionina. The hypoconulid is developed and located buccally. As for BPB 13, the size of the m/3 of BPB 30 enters into the range of variation of Pp. broomi (Tables 67; Fig. 13).

reMArks

In the literature, there is more information concerning the skulls than the mandibles to distinguish the different taxa of Papionini. So here, despite the preservation of BPB 1, this skull is most useful for the discussion. We must consider the biochronological age of the specimen and it is necessary to discuss older taxa (from East Africa) but also the Plio-Pleistocene Papionini. BPB 1 cannot belong to another Pliocene papionin taxon such as Pliopapio Frost, 2001 (5.2-4.2 Ma: Ethiopia) (Frost 2001; Jablonski & Frost 2010) which has a clear ante-orbital drop and an ophryonic groove. It cannot be attributed to Procercocebus antiquus either (Gilbert 2007a) which occurs at Taung (South Africa), and is characterized by an extremely straight nasal profile, widely divergent anterior temporal lines, relatively larger premolars, more prominent maxillary ridges and relatively more extensive maxillary/suborbital fossae than Parapapio. As indicated in the general description of BPB 1, the morphology of the interorbital and nasal regions indicates a slight concavity and lack of an ante-orbital drop which exclude the possibility that this specimen represents an individual of the smaller taxa of fossil Papio such as P. izodi and P. angusticeps [START_REF] Gilbert | -Papio cranium from the hominin-bearing site of Malapa: Implications for the evolution of the modern baboon cranial morphology and South African Plio-Pleistocene biochronology[END_REF][START_REF] Gilbert | Evolution of the modern baboon (Papio hamadryas): A reassessment of the African Plio-Pleistocene record[END_REF].

Three species of Parapapio are recognized in South Africa [START_REF] Jablonski N | Cenozoic Mammals of Africa[END_REF][START_REF] Harrison T | Cercopithecids (Cercopithecidae, Primates)[END_REF][START_REF] Gilbert | Cladistic analysis of extant and fossil African papionins using craniodental data[END_REF] during the Plio-Pleistocene: Pp. jonesi, Pp. broomi and Pp. whitei. Older fossils exist in the Early Pliocene at Langebaanweg [START_REF] Grine | Earliest primate remains from South Africa[END_REF] and Waypoint 160 (Gommery et al. 2008b(Gommery et al. , 2009[START_REF]Cercopithecoidea material from the Middle Pliocene site, Waypoint 160, Bolt's Farm, South Africa[END_REF] but they are too fragmentary or limited to assign to a specific taxon. In BPB 1, the maxillary ridge is more developed and the maxillary fossa is deeper than in Pp. jonesi. The premaxilla is not preserved in BPB 1 but it would not have extended much further anteriorly, as suggested by the anterior morphology of the maxilla, and as such the muzzle appears short and broad and its dorsum is flattened. All these features are typical of Pp. broomi. In Pp. whitei, the muzzle is long and narrow. The nasal bone and muzzle, especially in males, have a peaked aspect [START_REF] Gilbert | Cladistic analysis of extant and fossil African papionins using craniodental data[END_REF]. The morphology of BPB 1 is similar to Pp. broomi specimens such as BF 43 from Pit 23 at Bolt's Farm or M 202 from Makapansgat, so it can confidently be assigned to this species.

Pp. ado (Hopwood, 1936), a smaller species, is known from Kenya (Jablonski et al. 2008a) and Tanzania (Harrison 2011) (4.1-3.49 Ma). Its frontal and maxilla have similar morphologies [START_REF] Harrison T | Cercopithecids (Cercopithecidae, Primates)[END_REF] to Pp. broomi. These two species differ by the nasals which form a midline keel in Pp. broomi which is absent in Pp. ado. Despite the distortion of the nasals, the preserved transversal convexity in BPB 1 suggests that a midline keel was present. An older species exists, Pp. lothagamensis [START_REF] Ward C V | Cercopithecidae from Lothagam[END_REF][START_REF] Ward C V | Cercopithecidae from Lothagam[END_REF][START_REF] Jablonski N | Cenozoic Mammals of Africa[END_REF]Gilbert 2013) (7.4-c. 5 Ma: Kenya), but it is considered to be more primitive than the younger species [START_REF] Gilbert | Cladistic analysis of extant and fossil African papionins using craniodental data[END_REF].

The mandibles BPB 2 and BPB 3 probably belong to the same individual, which must have been a female. The teeth of these mandibles show a stage of wear corresponding to that of BPB 1. These two mandibles were found in a reduced sample of breccia taken from the "Parapapio spot". It is very likely that these three fossils belong to the same individual. 6,7); BPB 15 (Fig. 18D-F; Tables 6,7); BPB 27 (Fig. 18G-H). --Postcranial bones. BPB 5 (Fig. 20A,B); BPB 6 (Fig. 20C,D). desCription Isolated teeth BPB 14 (Fig. 18A-C; Tables 6,7). This isolated lower molar is probably a left m/2. The mesial contact surface is bigger and located more buccally than the distal one which is located more centrally. The enamel is thick and the buccal cusps are more worn than the lingual cusps. The protolophid and the hypolophid are low. The protoconid is the largest and the most deeply worn cusp. The metaconid is located slightly more mesially than the protoconid. The median clefts are narrow. The beginning of the mesial and distal roots is preserved. The anatomical features of this tooth correspond to the molars of papionines. BPB 14 is smaller than the m/2 of BPB 2 and BPB 3 but enters into the range variation of the m/2 of Pp. broomi (Tables 6,7; Fig. 12).

BPB 15 (Fig. 18D-F; Tables 6,7). The crown of this right p/4 is well preserved and is unworn. The roots are fragmentary. The protoconid and metaconid are close and clearly separated by a V-shaped transverse crest. These two cuspids are moderate salient from the crown and their apex is very slightly worn. The protoconid is slightly lower and is the most prominent cuspid. In buccal view (Fig. 18F), there is a slight mesio-buccal flange but also a deep distal and a mesial buccal cleft. The mesial fovea is small and moderately deep. The talonid is developed and high with a deep distal basin edged by a well-developed distal cingulum. The hypoconid is developed but narrower than the mesial part of the tooth. There is a small entoconid. The prehypocristid and the pre-entoconid are very short. There is a split on the lingual edge of the crown between the prehypocristid and the postprotocristid and between the pre-entoconid and the post-entoconid. The morphology of BPB 15 corresponds to a p/4 of papionines. The size of this premolar enters more into the range of variation of Pp. whitei than that of Pp. jonesi (Tables 6,7; Fig. 19).

BPB 27 (Fig. 18G,H). This small tooth should be a left di/1 and the apex of the root is broken (maximum height: 10.6 mm). The crown is very worn and presents an isosceles triangular outline (MD: 3 mm, BL: 3.9). The enamel is only preserved on the buccal surface (preserved height: 3.5 mm). The root is well preserved except for the apex (preserved height: 7.3 mm). The root is slightly curved. The buccal surface of the root is concave mesio-distally and the lingual one is slightly convex. The morphology of this incisor looks like some juvenile specimens of Plio-Pleistocene papionines.

Post-cranial bones. BPB 5 (Fig. 20A,B). This left patella is well preserved except for the disto-inferior part and the median side which have lost some superficial bone (maximum length: 18.2 mm; breadth: 12.9 mm; thickness: 8.3 mm). This patella presents a general morphology similar to WP 3 (a right patella from Waypoint 160, Bolt's Farm [Gommery et al. 2008b]), less elongated than in Papio ursinus. In anterior view (Fig. 20A), the medial part is more convex and shorter medio-laterally than the lateral part. In posterior view (Fig. 20B), the medial part of the articular facet is longer proximo-distally and shorter medio-laterally than the lateral part, and as well more concave proximo-distally. The distal apex is low, broad and has a rounded aspect with a groove posteriorly as in WP 3 (corresponding to the insertion of the lateral and medial Vasti and the M. rectus femoris). Despite the damage posteriorly, the proximal part of the patella is more elongated proximo-distally than the distal part. The morphology of this patella seems to suggest a mixed locomotor behaviour (terrestrial and arboreal).

BPB 6 (Fig. 20C,D). This fossil is a fragment of a right talus (length: 24.4 mm, breadth: 11 mm). Only a part of plantar surface is preserved with the head (Caput tali) and the neck (Collum tali), as well as the distal part of the body of the talus (Corpus tali) (Fig. 20C). The neck is broad relatively to its length (length 14.7 mm; breadth: 10.8 mm). On the head, the plantar part of the articular surface for the navicular is preserved and presents a slightly convex outline. The anterior and medial calcaneal articular surfaces are well preserved. These form a large proximo-distally elongated rectangle and occupy a large part of the surface along the medial edge of the talar neck. This morphology is typical of cercopithecines and differs from those of extant colobines where the two surfaces are distinct [START_REF] Olivier | -Les os du pied de semnopithèque[END_REF][START_REF] Strasser E | Pedal evidence for the origin and diversification of cercopithecid clades[END_REF][START_REF] Gilbert | -Miocene Cercopithecoidea from the Tugen Hills, Kenya[END_REF] but not in Paracolobus mutiwa [START_REF] Anderson | An assessment of the postcranial skeleton of the Paracolobus mutiwa (Primates: Colobinae) specimen KNM-WT 16827 from Lomekwi, West Turkana, Kenya[END_REF]. [START_REF] Gommery | Functional anatomy of the calcaneum and talus in Cercopithecinae (Mammalia, Primates, Cercopithecidae)[END_REF] have observed that the anterior calcaneal surface is more laterally developed in some cercopithecines such as Lophocebus and Miopithecus but not in Papio and many species of Macaca. In BPB 6, this surface is not developed laterally and is more similar to Papio. This morphology of the anterior and the medial articular surfaces is more related to weak rotation of the head of the talus as is usual in cercopithecines and different from the stronger rotation observed in colobines [START_REF] Strasser E | Pedal evidence for the origin and diversification of cercopithecid clades[END_REF]. Laterally, the fovea of the anterior astragalo-calcaneal ligament is moderately deep. The talar sulcus (Sulcus tali) is narrow and shallow laterally but broader and deeper medially below the groove for flexor hallucis longus.

The posterior calcaneal facet is preserved in its distal part. This facet seems deeply concave latero-distally to medio-proximally as in Papio. The junction with the groove for flexor hallucis longus is very salient. This groove has lost some superficial bone and is narrow. There is no swelling distally to the groove for the flexor hallucis longus as in Papio, Cercocebus and Cercopithecus (Strasser 1988;[START_REF] Gilbert | -Miocene Cercopithecoidea from the Tugen Hills, Kenya[END_REF].

BPB 6 is very damaged in dorsal view (Fig. 20D). Medially, a small fragment of the tibial malleolar facet is desCription Cranium BPB 7 (Fig. 21) General description and preservation. This specimen is the anterior part of the calvaria (maximum length: 54.9 mm; breadth: 83.5 mm) (Fig. 21A-C) comprised of two isolated pieces (Fig. 21D). It is less distorted than BPB 1. The left anterior part (corresponding to the upper part of the left orbit) is the most affected by distortion. Many cracks are present mainly in the posterior part. In anterior view, the upper part of the orbits, the glabella and the interorbital pillar are preserved. The latter is well preserved and broad (15.9 mm), typical of colobine monkeys, especially Cercopithecoides [START_REF] Freedman L | The fossil Cercopithecoidea of South Africa[END_REF][START_REF] Freedman L | Fossil and subfossil Primates from the limestone deposits at Taung, Bolt's Farm and Witkrans, South Africa[END_REF][START_REF]Evolutionary History of the Primates[END_REF][START_REF] Frost S. R. & Delson E | -Fossil Cercopithecidae from the Hadar Formation and surrounding areas of the Afar Depression, Ethiopia[END_REF]Jablonski et al. 2008b;[START_REF] Jablonski N | Cenozoic Mammals of Africa[END_REF]. This genus is common in South African Plio-Pleistocene sites. In superior view (Fig. 21A), there is an important ophryonic groove as is usual in Cercopithecoides.

There are a few bits of bone missing from the frontal but one of them, the biggest, presents an oval outline and probably corresponds to a predation mark.

Calvaria. In superior view (Fig. 21A), the ophryonic groove is deep and broad antero-posteriorly. Posterior to the ophryonic groove, the frontal has a rounded aspect. The temporal lines are well developed anteriorly with a strongly salient aspect as in BF 42A. Despite the damage, the postorbital constriction seems weak.

Orbital region. The superior orbital margins and parts of the lateral orbital margins are preserved and thick (thickness of the superior: 7.9 mm at left and 7.4 mm at right) (Fig. 21C). The supraorbital torus is well developed and thick. The preserved part suggests that the orbits were large. The lower aspect of the interorbital pillar suggests that the nasal and maxilla sutures are not fused; this frontal bone probably belongs to a sub-adult. The interorbital pillar is large. The glabellar region is moderately inflated anteriorly but this part is abraded. 2 Ma: Kenya), by its greater dimensions and thicker supraorbital torus despite the fact that this last feature seems variable in Cercopithecoides [START_REF] Anderson | Morphological diversity and species recognition in South African Cercopithecoides williamsi[END_REF][START_REF] Frost S. R | -Early Pliocene Cercopithecidae from Woranso-Mille (Central Afar, Ethiopia) and the origins of the Theropithecus oswaldi lineage[END_REF]. Some colobine remains from Woranso-Mille (Ethiopia) dated between 3.8 and 3.6 Ma were attributed to Cercopithecoides cf. meaveae [START_REF] Frost S. R. & Delson E | -Fossil Cercopithecidae from the Hadar Formation and surrounding areas of the Afar Depression, Ethiopia[END_REF][START_REF] Frost S. R | -Early Pliocene Cercopithecidae from Woranso-Mille (Central Afar, Ethiopia) and the origins of the Theropithecus oswaldi lineage[END_REF]. This species was known previously only at Leadu and Hadar (Ethiopia) desCription Mandible BPB 8 (Fig. 22A,B). This fossil (maximum length: 20.8 mm; height: 20 mm) is the distal part of the ascending ramus and the mandibular condyle of a right mandible. The median part of the condyle is damaged and the lateral part is very salient (length: 9.3 mm; breadth: 16.6 mm).

Remarks

BPB 26 (Fig. 22C,D). The preserved part (maximum length: 26.1 mm; height: 24.7 mm) comprises the ascending ramus and the mandibular condyle (maximum length: 7.8 mm; preserved breadth: 6.4 mm) of a right mandible. 1994). The proximal extremity is slightly larger (preserved breadth: 6.9 mm, height: 5.4 mm) than the distal extremity (breadth: 6.1 mm, height: 4.1 mm). The proximal articular facet presents a double concavity as is usual in a median phalanx. In dorsal view (Fig. 22E), a small hole is present in the distal extremity and corresponds probably to a predator mark. In volar view (Fig. 22F), along all the shaft, there are extensions underlined by deep depressions which correspond to the insertion of collateral ligaments of the manual interphalangeal joint. The central part of the shaft is convex medio-laterally and forms a pillar. The proximal extremity presents a smooth bulge on each side. There is a small oval cavity which possibly corresponds to a mark of predation.

The distal extremity presents a trochlear aspect. In plantar view, the medial condyle of this trochlea is the narrower and the more convex of the two condyles. It is not possible to say more about this phalanx. et al. 2022). This species is also present at Makapansgat Limeworks in Member 3 [START_REF] Freedman L | The fossil Cercopithecoidea of South Africa[END_REF][START_REF] Freedman L | Some news fossil, Cercopithecoid specimens from Makapansgat, South Africa[END_REF][START_REF] Freedman L | South African fossil Cercopithecoidea: A re-assessment including a description of new material from Makapansgat, Sterkfontein and Taung[END_REF][START_REF] Maier | New fossil Cercopithecoidea from the lower Pleistocene cave deposits of the Makapansgat Limeworks, South Africa[END_REF] South Africa during the mid-Pliocene. We conclude that primate migrations between East and Southern Africa occurred earlier than previously thought. Southern Africa could also have been an important centre of primate diversification during the Pliocene and the beginning of the Quaternary. Although the remains of Early Pliocene Cercopithecoidea from Southern Africa are scarce and fragmentary [START_REF] Grine | Earliest primate remains from South Africa[END_REF]Gommery et al. 2008bGommery et al. , 2009[START_REF]Cercopithecoidea material from the Middle Pliocene site, Waypoint 160, Bolt's Farm, South Africa[END_REF][START_REF] Brink | Waypoint 160, Bolt's Farm Cave System: First in situ primate remains[END_REF], we cannot rule out the possibility that some Cercopithecoidea evolved first in Southern Africa before dispersing to East Africa. The Pliocene is recognized as a globally warm period with permanent El Niño conditions and there were wet conditions in East Africa particularly during the Early Pliocene [START_REF] Jablonski N | Cenozoic Mammals of Africa[END_REF][START_REF]African climate change and faunal evolution during the Pliocene-Pleistocene[END_REF][START_REF] Feakins S | Global and African regional climate during the Cenezoic[END_REF]. A progressive cooling starting at around 3.2 Ma [START_REF] Jablonski N | Cenozoic Mammals of Africa[END_REF][START_REF]African climate change and faunal evolution during the Pliocene-Pleistocene[END_REF][START_REF] Feakins S | Global and African regional climate during the Cenezoic[END_REF] led to a gradual aridification of the environment in East Africa but also in South Africa. The data concerning the climatic and environmental changes in Southern Africa are limited for the Pliocene in comparison with East Africa. The palaeoenvironment at Waypoint 160 in the Early Pliocene seems to have been dry and open as suggested by the rodent fauna and especially the presence of Petromus (Sénégas 2004) and Euryotomys (Sénégas 2001). BPB is one of the two potential known sites in Southern Africa to be close to the Early/Late Pliocene boundary. The environment at BPB was probably not as open and dry as at Waypoint 160 but it was probably a grassland-woodland mosaic as suggested by the rodents (under study) and the bald ibis (Geronticus thackerayi) [START_REF] Pavia | Geronticus thackerayi, sp. nov. (Aves, Threskiornithidae), a new ibis from the hominin-bearing locality of Kromdraai (Cradle of Humankind, Gauteng, South Africa)[END_REF]Pavia et al. 2017). The presence of suids as Gerontochoerus koobiforaensis (Pickford et al. 2019) at BPA seems to confirm that. It is the first known report of this environment for the Pliocene in this part of South Africa (Sterkfontein Valley, Gauteng Province). This environment seems to allow the development of Parapapio. It is probably the same for Cercopithecoides, even if it presents anatomical characteristics showing that it spent a good part of the time on the ground, like some to modern large-bodied colobines it probably climbed trees to feed and to protect itself from ground-dwelling predators (Jablonski & Frost 2010).

In the Southern Hemisphere, some cooler events related to the Benguela upwelling system [START_REF] Marlow | Upwelling intensification as part of the Pliocene-Pleistocene climate transition[END_REF] in the Atlantic are observed during the mid-Pliocene. In the Antarctic [START_REF] Lucchi R | Circum-Antarctic warming events between 4 and 3.5 Ma recorded in marine sediments from the Prydz Bay (ODP Leg 188) and the Antarctic Peninsula (ODP Leg 178) margins[END_REF], the cooling transition started earlier, at 3.7-3.6 Ma. These events could have possibly reduced the effect of the permanent El Niño in Southern Africa earlier than in East Africa. At present it is impossible to confirm but something caused a transitional phase which resulted in less dry conditions, necessary for a grassland-woodland mosaic, before the gradual aridification of the environment known since the beginning of the Early Pleistocene in the Sterkfontein Valley. These environmental conditions were probably very favourable to the expansion of certain primate taxa from East Africa to southern Africa and/or a diversification into southern Africa of taxa already present. Further research at both BPB and BPA but also at Waypoint 160 may provide information to answer these questions. [START_REF] Freedman L | The fossil Cercopithecoidea of South Africa[END_REF][START_REF] Freedman L | Fossil and subfossil Primates from the limestone deposits at Taung, Bolt's Farm and Witkrans, South Africa[END_REF][START_REF] Frost S. R | -New Early Pliocene Cercopithecidae (Mammalia: Primates) from Aramis, Middle Awash Valley, Ethiopia[END_REF][START_REF] Frost S. R. & Delson E | -Fossil Cercopithecidae from the Hadar Formation and surrounding areas of the Afar Depression, Ethiopia[END_REF] this study) used for the Figure 19.

  located in the North of South Africa. The major part of these GEODIVERSITAS • 2023 • 45 (5) collections come from a restricted area in the western part of Gauteng Province and the neighbouring part of the North-West Province. Recognized as the Cradle of Humankind World Heritage Site (COH WHS) in 1999 by the UNESCO and is mainly located around the Sterkfontein Valley. Robert Broom was the first to conduct regular research in this area from 1936 to his death in 1951, in particular on the Palaeokarst System of Bolt's Farm (called Bolt's Farm in this article) (Thackeray et al. 2008).

  fig. 1. -Map of the locality of Brad Pit (BP). The different loci: BPA, Brad Pit A; BPB, Brad Pit B; BPE, Brad Pit Extension (newly discovered). "Parapapio spot" is indicated by a black star.

  fig. 2. -View of the locus BPB with the "Parapapio spot" in the white circle.

  rich and is dominated by a new taxon of rodent. This taxon would represent the last species of Euryotomys Pocock, 1976, in the fossil record prior to the occurrence of Otomys F. Cuvier, 1824, which appears in younger Late Pliocene and Early Pleistocene sites. The oldest occurrence of Otomys is known at Makapansgat Limeworks (Pocock 1987), more specifically at Rodent Corner probably dated to 3.5-3.3 Ma (Hopley et al. 2006, 2007) and in Member 3 dated to 3.03-2.61 Ma (Reed et al. 2022, modified from Herries et al. 2013). So far no Euryotomys remains have been found in Members 2 and 4 of Sterkfontein but some Otomys (Pocock 1987; Avery 2001; Reynolds & Kibii 2011) have been recovered. The Member 2 is dated to approximately 3.67 Ma (Granger et al. 2015; Van Couvering & Delson 2020). The Member 4 was previously dated between 2.8 and 2 Ma by different methods (Pickering & Kramers 2010; Herries & Shaw 2011; Reynolds & Kibii 2011; Van Couvering & Delson 2020) and current dating techniques date the deposit at 3.4 Ma (Granger et al. 2022). The new species of Euryotomys, more derived than E. bolti Sénégas & Avery, 1998, from Waypoint 160, would place the site biochronologically between Waypoint 160 and the oldest deposits at Makapansgat suggesting a middle Pliocene age for BPB. This new species of Euryotomys is also present at BPA. Some teeth of a Suidae belonging to Gerontochoerus koobiforaensis Pickford, 2013, confirm the Pliocene age of the infill, probably middle Pliocene (Pickford et al. 2019).

  fig. 3. -Specimen BPB 1 (Parapapio broomi Jones, 1937) still embedded in the in situ breccia before extraction: muzzle extremity (A); skull vault (B); skull back (C); right M3/ (D).

  Gommery D. et al.

  1 mm. The BPB fossils have been compared with some Parapapio and Cercopithecoides specimens and as well other Papionina in the collections from Sterkfontein (from Member 4) and Taung (from Pit 5, fossils collected by the Southern Section of the University of California African Expedition (UCAE) led by Charles L. Camp between 1947-1948) curated in the Ditsong National Museum of Natural History (DNMNH) at Pretoria and from Makapansgat Limeworks (Members 3 & 4) and Pit 23 of Bolt's Farm Cave System (as well fossils collected by the UCAE) in the University of the Witwatersrand (WU) in Johannesburg (Table

  fig. 4. -Preliminary observations concerning the breccia infilling of the "Parapapio spot" at the locus BPB: A, synthetic log of the locus BPB showing the stratigraphic position of Parapapio fossils within Unit 2; B, geological details of the "Parapapio spot" evidencing that the original bedding has been partly deformed by soft sediment deformation features at its topmost part.

  fig. 5. -Skull of Parapapio broomi Jones, 1937, specimen BPB 1: A, fronto-right lateral view (as preserved naturally); B, frontal view; C, right lateral view (of face); D, dorsal view; E, inferior view. Scale bar: 10 mm.

  fig. 6. -Teeth of Parapapio broomi Jones, 1937, specimen BPB 1: A, occlusal view of the right tooth row P4/-M3/; B, oblique-lingual view of M2/ and M3/. Scale bar: 10 mm. table 5. -Measurements of the mesio-distal length in mm of the dental segments (from Freedman 1957, 1965; this study) (the quantity of specimens is indicated between the brackets).

  ) unlike C. williamsi which presents a well-developed one. The buccal surface of the posterior part of the mandibular corpus

  fig. 11. -Right mandible of Parapapio broomi Jones, 1937, specimen BPB 2: A, buccal view; B, lingual view; C, superior view; D, occlusal view of m/1-m/3. Scale bar: 10 mm.

  fig. 14. -Left mandible of Parapapio broomi Jones, 1937, specimen BPB 3: A, buccal view; B, lingual view; C, superior view; D, occlusal view of p/4-m/3. Scale bar: 10 mm.

  fig. 16. -Distal part of the left mandible of Parapapio broomi Jones, 1937, specimen BPB 13: A, buccal view; B, lingual view; C, superior view; D, occlusal view of m/3. Scale bar: 10 mm.

  Tribe pApionini Burnett, 1828 Subtribe pApioninA Burnett, 1828 Papionina indet. MAteriAl exAMined. -Isolated teeth. BPB 14 (Fig. 18A-C; Tables

  fig. 17. -Part of the left mandible of Parapapio broomi Jones, 1937, specimen BPB 30: A, buccal view; B, lingual view; C, occlusal view of m/3. Scale bar: 10 mm.

  fig. 18. -A-C, Left lower molar of Papionina indet., specimen BPB 14: A, buccal view; B, lingual view; C, occlusal view; D-F, right lower fourth premolar (p/4) of Papionina indet., specimen BPB 15: D, buccal view; E, lingual view; F, occlusal view; G, H, lower left deciduous first incisor (di/1) of Papionina indet., specimen BPB 27: G, labial view; H, lingual view. Scale bar: 10 mm.

  fig.19. -Bivariate plots MD × BL (mesio-distal length × maximum bucco-lingual breadth in mm) of lower fourth premolars (p/4) of different taxa of fossil Papionines from South Africa with some from East Africa (data from[START_REF] Freedman L | The fossil Cercopithecoidea of South Africa[END_REF][START_REF] Freedman L | Fossil and subfossil Primates from the limestone deposits at Taung, Bolt's Farm and Witkrans, South Africa[END_REF][START_REF] Frost S. R | -New Early Pliocene Cercopithecidae (Mammalia: Primates) from Aramis, Middle Awash Valley, Ethiopia[END_REF][START_REF] Frost S. R. & Delson E | -Fossil Cercopithecidae from the Hadar Formation and surrounding areas of the Afar Depression, Ethiopia[END_REF] this study) (Appendix 7).

.

  The morphology and the size of the specimen are comparable to the homologous frontal areas of different specimens of the very large colobine Cercopithecoides williamsi such as BF 42A (Pit 23, BFCS) or AD 1326-6/M 236 (type specimen), M 2999/MP 113 and M 3055 (from Makapansgat Limeworks). In the absence of a complete muzzle and dentition[START_REF] Frost S. R. & Delson E | -Fossil Cercopithecidae from the Hadar Formation and surrounding areas of the Afar Depression, Ethiopia[END_REF][START_REF] Jablonski N | Cenozoic Mammals of Africa[END_REF], we provisionally assign the fossil BPB 7 to C. cf. williamsi. It differs from the older species, C. kerioensis[START_REF] Ward C V | Cercopithecidae from Lothagam[END_REF][START_REF] Ward C V | Cercopithecidae from Lothagam[END_REF] Jablonski & Frost 2010) (c. 5-4.

  between 3.4 to 3.28 Ma (Frost & Delson 2002; Jablonski & Frost 2010). The remains from Woranso-Mille are larger than the youngest remains from Leadu and Hadar but are still smaller than those of C. willamsi. Another large colobine, similar in size to the genus Paracolobus Leakey, 1969, is presumed to occur at Makapansgat (Jablonski & Frost 2010) but it is represented by isolated teeth. Paracolobus differs from Cercopithecoides by a marked postorbital constriction, not present on BPB 7. Family CerCopitheCidAe Gray, 1821 Cercopithecidae indet. MAteriAl exAMined. -Mandible. BPB 8 (Fig. 22A, B); BPB 26 (Fig. 22C, D). -Postcranial bone. BPB 19 (Fig. 22E, F).

  fig. 21. -Anterior part of calvaria of Cercopithecoides cf. williamsi Mollett, 1947, specimen BPB 7: A, superior view; B, inferior view; C, frontal view; D, two isolated fragments of the calvaria. Scale bar: 10 mm.

  DISCUSSION AND CONCLUSION It is not uncommon to find Cercopithecoides and Parapapio associated in South African Plio-Pleistocene sites. At BPB, we have potentially the oldest known co-occurrence of these two taxa, the second oldest could be at Sterkfontein-Member 2 (Pickering et al. 2004; Reynolds & Kibii 2011). The BPB Pp. broomi specimen indicates that this species constitutes an old South African taxon. It is almost contemporary with the oldest known remains of Pp. jonesi surely known in the Lower Kada Hadar member of the Hadar Formation in Ethiopia (3.18-2.92 Ma) and perhaps at Ahmado (3.22-3.18 Ma) also in Ethiopia (Frost & Delson 2002; Jablonski & Frost 2010) which represents the earliest papionin with a nearly Pan-African distribution but which cannot represent the base of Parapapio in South Africa. At Koobi Fora in Kenya, except for an isolated m/3 dated at around 3.5 Ma, most of the material of C. williamsi was considered to be younger (2.5-1.5 Ma) (Jablonski et al. 2008b). This species is known at Sterkfontein (Pickering et al. 2004; Reynolds & Kibii 2011) from Member 2 dating approximately 3.67 Ma (Granger et al. 2015; Van Couvering & Delson 2020) and Member 4 dated recently around 3.4 Ma (Granger

  fig. 22. -A, B, Distal part of the ascending ramus of a right mandible of Cercopithecidae indet., specimen BPB 8: A, lateral view; B, distal view; C, D, distal part of the ascending ramus of a right mandible of Cercopithecidae indet., specimen BPB 26: C, lateral view; D, medial view; E, F, right intermediate manual phalanx of Cercopithecidae indet., specimen BPB 19: E, dorsal view; F, volar view. Scale bar: 10 mm.

table 1 .

 1 -List of the specimens by taxon, site and anatomical part of the different taxa of original fossil primates from South Africa used for anatomical comparison.

table 3 .

 3 -Measurements in mm of upper teeth of BPB 1.

	BPB1	P4/	M1/	M2/	M3/
	MD	6.6	9.3	11.4	12.4
	BL	8.9	-	-	-
	Mb	-	9.7	11	11.4
	Dd	-	8.9	9.8	9.3
	Dth	-	-	-	5.1

GEODIVERSITAS • 2023 • 45 (5)

table 7 .

 7 -Synthesis of variation in lower tooth measurements in mm (MD, mesio-distal length; BL, maximum bucco-lingual breadth) in different taxa of fossil Papionines from South Africa with some from East Africa from the literature (from[START_REF] Freedman L | The fossil Cercopithecoidea of South Africa[END_REF][START_REF] Freedman L | Fossil and subfossil Primates from the limestone deposits at Taung, Bolt's Farm and Witkrans, South Africa[END_REF][START_REF] Frost S. R | -New Early Pliocene Cercopithecidae (Mammalia: Primates) from Aramis, Middle Awash Valley, Ethiopia[END_REF][START_REF] Frost S. R. & Delson E | -Fossil Cercopithecidae from the Hadar Formation and surrounding areas of the Afar Depression, Ethiopia[END_REF] and this study (the quantity of specimens is indicated between the brackets).

		p/4		m/1	m/2		m/3
	Taxon	MD	BL	MD	BL	MD	BL	MD	BL
	Pp. broomi BF	6.6 (1)	6.9 (1)	9.1 (1)	7.9 (1)	11 (1)	9.5 (1)	13.5-14.2 (2)	9.3-9.9 (2)
	Pp. broomi Sts	5.7-8.3 (16)	5.9-7.7 (16) 7.4-10.3 (14) 7.1-8.6 (14) 9.5-12.4 (16) 8.8-10.8 (16) 11.5-15.6 (16) 8.7-10.7 (16)
	Pp. broomi M	6.4-7.4 (3)	6.6-7.9 (3)	8.6-9.6 (5)	7.8-8.6 (5)	10.5-11.8 (5) 9.6-10.4 (5)	14-15.3 (3) 10.3-10.5 (3)
	Pp. whitei Sts	6.9-8.1 (7)	6.3-7.7 (7)	9.1-10.2 (7)	7.4-9.1 (7)	11.5-13.1 (6)	9-10.9 (6)	15.9-16.5 (5) 10.6-11.6 (5)
	Pp. whitei M	7-7.2 (2)	7.4-8.1 (2)	10.9 (1)	8.4 (1)	-	-	16.5 (1)	11.6 (1)
	Pp. jonesi Sts	6-7.5 (18)	5.9-7.5 (18)	7.8-9.7 (11)	6.8-7.8 (11) 9.6-11.4 (14) 8.6-10.1 (14) 12.2-13.7 (10) 8.5-10 (10)
	Pp. jonesi M	-	-	8.3-9 (3)	6.7-7.3 (3)	9.5-10.8 (2)	7.6-9 (2)	11.1-13.2 (4)	7.9-9.3 (4)
	Pp. jonesi EA	6.3-6.4 (2)	6-6.5 (2)	7.9-10 (2)	7.2-7.6 (2)	10-10.4 (2)	9.2 (2)	13.6-14.1 (4)	8.2-9.7 (4)
	Pr. antiquus T	7.5 (1)	7.1 (1)	9.8-10.3 (2)	7.8-8.3 (2)	12.1 (1)	9.6 (1)	14.6 (1)	10 (1)
	P. angusticeps KA 7.7-8.3 (4)	6.4-7.4 (4)	7.8-8.7 (4)	9.4-10.4 (4) 11.6-12.6 (5) 9.3-10.1 (5)	14.2 (7)	9.9-10.7 (7)

GEODIVERSITAS • 2023 • 45 (5)

  -Mesio-distal length (MD) and maximum bucco-lingual breadth (BL) in mm of upper fourth premolars (P4/) of different taxa of fossil Papionines from South Africa with some from East Africa (data from[START_REF] Freedman L | The fossil Cercopithecoidea of South Africa[END_REF][START_REF] Freedman L | Fossil and subfossil Primates from the limestone deposits at Taung, Bolt's Farm and Witkrans, South Africa[END_REF][START_REF] Frost S. R | -New Early Pliocene Cercopithecidae (Mammalia: Primates) from Aramis, Middle Awash Valley, Ethiopia[END_REF][START_REF] Frost S. R. & Delson E | -Fossil Cercopithecidae from the Hadar Formation and surrounding areas of the Afar Depression, Ethiopia[END_REF] this study) used for the Figure7. -Mesio-distal length (MD) and maximum bucco-lingual breadth (BL) in mm of upper first molars (M1/) of different taxa of fossil Papionines from South Africa with some from East Africa (data from[START_REF] Freedman L | The fossil Cercopithecoidea of South Africa[END_REF][START_REF] Freedman L | Fossil and subfossil Primates from the limestone deposits at Taung, Bolt's Farm and Witkrans, South Africa[END_REF][START_REF] Frost S. R | -New Early Pliocene Cercopithecidae (Mammalia: Primates) from Aramis, Middle Awash Valley, Ethiopia[END_REF][START_REF] Frost S. R. & Delson E | -Fossil Cercopithecidae from the Hadar Formation and surrounding areas of the Afar Depression, Ethiopia[END_REF] this study) used for the Figure8. -Mesio-distal length (MD) and maximum bucco-lingual breadth (BL) in mm of upper second molars (M2/) of different taxa of fossil Papionines from South Africa with some from East Africa (data from[START_REF] Freedman L | The fossil Cercopithecoidea of South Africa[END_REF][START_REF] Freedman L | Fossil and subfossil Primates from the limestone deposits at Taung, Bolt's Farm and Witkrans, South Africa[END_REF][START_REF] Frost S. R | -New Early Pliocene Cercopithecidae (Mammalia: Primates) from Aramis, Middle Awash Valley, Ethiopia[END_REF][START_REF] Frost S. R. & Delson E | -Fossil Cercopithecidae from the Hadar Formation and surrounding areas of the Afar Depression, Ethiopia[END_REF] this study) used for the Figure9.Gommery D. et al. -Mesio-distal length (MD) and maximum bucco-lingual breadth (BL) in mm of upper third molars (M3/) of different taxa of fossil Papionines from South Africa with some from East Africa (data from[START_REF] Freedman L | The fossil Cercopithecoidea of South Africa[END_REF][START_REF] Freedman L | Fossil and subfossil Primates from the limestone deposits at Taung, Bolt's Farm and Witkrans, South Africa[END_REF][START_REF] Frost S. R | -New Early Pliocene Cercopithecidae (Mammalia: Primates) from Aramis, Middle Awash Valley, Ethiopia[END_REF][START_REF] Frost S. R. & Delson E | -Fossil Cercopithecidae from the Hadar Formation and surrounding areas of the Afar Depression, Ethiopia[END_REF] this study) used for the Figure10. appenDix 5. -Mesio-distal length (MD) and maximum bucco-lingual breadth (BL) in mm of lower second molars (m/2) of different taxa of fossil Papionines from South Africa with some from East Africa (data from[START_REF] Freedman L | The fossil Cercopithecoidea of South Africa[END_REF][START_REF] Freedman L | Fossil and subfossil Primates from the limestone deposits at Taung, Bolt's Farm and Witkrans, South Africa[END_REF][START_REF] Frost S. R | -New Early Pliocene Cercopithecidae (Mammalia: Primates) from Aramis, Middle Awash Valley, Ethiopia[END_REF][START_REF] Frost S. R. & Delson E | -Fossil Cercopithecidae from the Hadar Formation and surrounding areas of the Afar Depression, Ethiopia[END_REF] this study) used for the Figure12.Gommery D. et al. -Mesio-distal length (MD) and maximum bucco-lingual breadth (BL) in mm of lower third molars (m/3) of different taxa of fossil Papionines from South Africa with some from East Africa (data from[START_REF] Freedman L | The fossil Cercopithecoidea of South Africa[END_REF][START_REF] Freedman L | Fossil and subfossil Primates from the limestone deposits at Taung, Bolt's Farm and Witkrans, South Africa[END_REF][START_REF] Frost S. R | -New Early Pliocene Cercopithecidae (Mammalia: Primates) from Aramis, Middle Awash Valley, Ethiopia[END_REF][START_REF] Frost S. R. & Delson E | -Fossil Cercopithecidae from the Hadar Formation and surrounding areas of the Afar Depression, Ethiopia[END_REF] this study) used for the Figure13. appenDix 7. -Mesio-distal length (MD) and maximum bucco-lingual breadth (BL) in mm of lower fourth premolars (p/4) of different taxa of fossil Papionines from South Africa with some from East Africa (data from Freedman

	GEODIVERSITAS • 2023 • 45 (5) Locality Parapapio broomi Jones, 1937 MD Sterkfontein 6.7 6.7 6.2 6.5 6.3 6.5 5.8 6.5 6.5 6.2 6.2 5.4 6.7 6.9 6.1 6.1 Makapansgat 6.5 6.6 6.3 6.6 6.8 6.4 6.3 Parapapio whitei Broom, 1940 Sterkfontein 6 7.8 7 6.5 7 7.1 7.4 6.7 6.2 Makapansgat 6 6.1 6.1 7 appenDix 1. Gommery D. et al. Locality MD Parapapio broomi Jones, 1937 Bolt's Farm 9.9 Sterkfontein 9.8 9.8 9.8 8.4 9.2 9.9 9.8 9.8 10 10.1 8.8 9.7 9 9.6 9.5 Makapansgat 9.2 9.9 10.2 10.3 9.8 9 Parapapio whitei Broom, 1940 Sterkfontein 9.7 10.7 9.5 10.6 11 10.9 10.1 Locality MD Parapapio broomi Jones, 1937 Bolt's Farm 11.2 10.8 11.5 11.4 Parapapio whitei Broom, 1940 Sterkfontein 11.8 13.2 11 13 13.4 13.2 13 11.8 11.8 Makapansgat 10.9 11.1 11.1 11.4 11.4 10.8 appenDix 3. Locality MD Parapapio broomi Jones, 1937 BPA 13.2 Parapapio whitei Broom, 1940 Sterkfontein 12.4 12.8 11.6 Makapansgat 10.4 10.2 11.4 12.1 appenDix 4. Locality MD Parapapio broomi Jones, 1937 Bolt's Farm 11 Locality MD Parapapio broomi Jones, 1937 Bolt's Farm 13.5 appenDix 6. Locality MD Parapapio broomi Jones, 1937 Bolt's Farm 6.6 appenDix 2. GEODIVERSITAS • 2023 • 45 (5) 11.7 Sterkfontein 10.8 11.9 10.3 11.2 10.5 10 11.1 9.8 11.6 10.8 11 10.8 10.7 10.9 10.9 10.8 11.2 11.2 10.5 10.6 11.2 10.8 Makapansgat 10.2 11.5 11 10.7 11 12.5 11.4 11.1 10.7 11 10.8 11 10.8 10.6 Makapansgat 9.5 7.2 Bolt's Farm 11.7 10.9 Sterkfontein 11.4 11.6 10.8 10.3 11.3 10.9 11.1 11.1 10.6 10.7 10.1 10.8 11.1 11 10.5 10.5 10.3 11.5 10 10.6 Makapansgat 10.8 11 10.6 11.2 10.6 10.2 11.2 Sterkfontein 10.5 11.5 10.7 10.9 10.5 9.6 11.3 12.4 10.9 12.4 9.9 9.5 12.2 10.5 11.7 11.2 Makapansgat 10.5 11.5 11.3 11.8 11.8 Parapapio whitei Broom, 1940 14.2 Sterkfontein 11.9 12.9 13.5 13.4 13.2 14.9 14.2 14.6 11.5 12.1 11.9 14.6 14.6 12.5 15.6 15.3 Sterkfontein 6.1 6.8 6.5 8.3 8.2 6.2 7.5 6.7 7.5 6.8 5.7 7.2 6.8 7.1 6.9 6.7 Makapansgat 15.3 15.3 14 Parapapio whitei Broom, 1940 Sterkfontein 16.5 Makapansgat 6.4 7.4 7.3 Parapapio whitei Broom, 1940 Sterkfontein 7.6 7.4 Sterkfontein 12.1 12.7 13.1 11.7 11.5 12.5 Makapansgat 7 15.3 16.5 15.5 16.5 Makapansgat 16.5 7.7 8.1 7.2 6.9 7.4	BL 8.4 8.6 7.3 7.1 8.5 8.1 7.9 7.1 8.7 8.3 8.5 7.8 8.7 9.6 8.4 7.7 8.3 8.5 8.4 9 9 8.5 8.5 7.8 9.1 7.9 9.2 8.6 8.6 8.7 8.6 8.2 8.7 9 9.5 9.1 BL 9.9 9.6 9.9 9.6 8.7 9.8 9.5 9.6 9.6 9.3 9.6 9.4 9.5 9 9.9 9 9.3 9.4 9.5 9.3 10.1 9.6 9.8 10.5 9.1 10.6 10.9 11 9.5 BL 11.7 11 11.8 11 12.3 12.6 10.3 12.1 12.5 13 13 13.2 11.7 10.6 10.9 12.6 12.8 11.7 11 BL 11 12.6 11.8 12.5 10.4 10.4 12.2 11.6 BL 9.5 BL 9.9 BL 6.9 11.7 11.1 12 10.2 10.5 11.8 10.3 11.7 10.9 10.9 10.3 10.5 12.1 10.9 10.9 10.9 10.1 11.4 10.6 10.7 10.1 11 10.5 10.8 11 10.9 11 12.1 12.3 11.7 11.8 12.2 10.5 10.8 10.3 9 11.6 7.6 7.4 10.8 10.5 11.2 11 11.1 10.4 10.5 10 10 10.6 11.4 11.5 10.4 10.8 10 10.1 10.8 10.1 10.2 12.3 9.9 10.3 10.4 12 12.1 11.7 11.5 11.2 11.8 8.8 9.3 8.9 8.8 9.4 8.8 10.6 9.7 10.6 10.4 8.9 9.2 9.9 9.1 10.8 10.4 10 10.1 10.4 9.6 10 9.3 8.9 9.2 9.5 9.3 11 10.2 9.9 10.2 8.7 9.1 8.8 10.3 10 9.1 10.7 10.1 6.5 6 5.9 7.4 7.2 6.7 7.2 6.9 6.6 7.1 6.1 6.9 6.5 6.6 6.6 7.7 10.5 10.5 10.3 11.1 6.6 7.9 7.3 7.7 6.9 10.9 10.4 10.5 10.1 9 10.5 8.1 10.6 10.8 11.1 10.8 11.6 6.3 6.9 6.5 7.1 7.2	APPENDICES Specimen Sts 251 Sts 260 Sts 261 Sts 264 Sts 267 Sts 297 Sts 368A Sts 377 Sts 383B Sts 385A Sts 390 Sts 398A Sts 414B Sts 424 Sts 539 Sts 544 MP2 (M202) MP 224 MP 224 M205A,B M2963 M2964 M2966 Sts 259 Sts 263 Sts 266 Sts 336 Sts 343 Sts 370A Sts 370 B Sts 389B Sts 548 MP 151 MP 76 MP 76 M2973 Locality Parapapio jonesi Broom, 1940 MD Sterkfontein 5.6 5.7 5.8 6.1 5.2 5.2 5.6 East Africa 5.4 5.8 Procercocebus antiquus (Haughton, 1925) Taung 7.4 7.3 Papio izodi Gear, 1926 Taung 7.6 Papio angusticeps (Broom, 1940) Kromdaai A 7.9 8.5 7.9 7.7 8.7 7.8 Cooper's 7.6 8.3 8.1 8.4 Specimen BF 43 Sts 251 Sts 253 Sts 260 Sts 264 Sts 267 Sts 274 Sts 297 Sts 301 Sts 377 Sts 383B Sts 385 Sts 390 Sts 414B Sts 539 Sts 544 MP 2 (M202) MP 2 (M202) MP 224 MP 224 M2963 M3037 Sts 259 Sts 263 Sts 266 Sts 336 Sts 343 Sts 370A Sts 548 Locality MD BL BL 7.8 7.5 7.5 6.1 7.3 7 7.4 7 9.5 8 7.5 7.5 6 7.3 7.4 6.1 7 6.7 7.6 6.4 6.9 7.2 Makapansgat 9.5 10.2 9.6 10.3 8.7 8.8 Parapapio jonesi Broom, 1940 Sterkfontein 7.3 9 8.9 9 8.9 8.7 7.6 9 Makapansgat 9.1 8.9 Taung 8.4 8.3 East Africa 8 8.2 8.2 10.2 Procercocebus antiquus (Haughton, 1925) Taung 10 9 Papio izodi Gear, 1926 Taung 10.5 10 9.7 9.5 10.8 10.8 Papio angusticeps (Broom, 1940) Kromdaai A 10.5 10.8 9.6 10.2 8.8 10.6 8.9 9 Cooper's 8.7 10.3 9.2 9.5 9.9 10.3 Specimen BF 56786 M3011 M3015 M3037 Sts 259 Sts 263 Sts 266 Sts 336 Sts 370A Sts 370 B Sts 462 Sts 503 Sts 548 MP 151 MP 151 MP 76 MP 76 M2963 M3011 Locality MD BL Parapapio jonesi Broom, 1940 Sterkfontein 9.5 10.4 Specimen BPA230 Sts 259 Sts 303 Sts 503 MP 151 MP 151 MP 76 M2962 Locality MD BL Parapapio jonesi Broom, 1940 Sterkfontein 10.9 10.4 Specimen BF 56796 Locality MD BL Parapapio jonesi Broom, 1940 Sterkfontein 10.4 10.1 Specimen BF 56796 Locality MD BL Parapapio jonesi Broom, 1940 Sterkfontein 12.7 8.5 Specimen BF 56796 Locality MD BL Parapapio jonesi Broom, 1940 Sterkfontein 7 6.6 BF 43 Sts 251 Sts 260 Sts 261 Sts 264 Sts 267 Sts 274 Sts 297 Sts 325 Sts 354 Sts 377 Sts 378 A-D Sts 379A Sts 380B Sts 380B Sts 383B Sts 385A Sts 388A Sts 397 Sts 398A Sts 410A Sts 414B Sts 544 MP 2 (M202) MP 2 (M202) MP 224 MP 224 M205 M 635 M2969 10.9 11 M3037 M2966 11.6 12.4 M3002 M2992 M2964 10.9 12.6 M2977 M215 M3062 M2963 10.3 10.3 10.5 10.1 9.8 9.7 9.8 10.1 10 9.8 9.2 9.4 9.2 10.4 Makapansgat 9.5 10.2 East Africa 10.6 11.9 Procercocebus antiquus (Haughton, 1925) Taung 11.8 10.6 11.8 11.6 11.9 10.7 11.6 11 Papio izodi Gear, 1926 Taung 12.8 11.8 11.8 11.2 11.8 11.5 Papio angusticeps (Broom, 1940) Kromdaai A 11.8 11.6 11.5 11.7 10.4 11.2 10.8 12.6 Cooper's 10.8 12.2 11 12.4 12.4 12.7 BF 43 BF 56786 Sts 261 Sts 261 Sts 267 Sts 325 Sts 378A Sts 380B Sts 383B Sts 385 Sts 388A Sts 388B Sts 390 Sts 393 Sts 397 Sts 397 Sts 398A Sts 410A Sts 414B Sts 534 Sts 535 Sts 544 MP2 (M202) M 205 M 635 M2963 M2964 M2970 M2975 9.8 10.3 9.7 9.9 9.5 10.6 9.6 10.6 9.7 9.9 9.6 9.8 8.9 9.3 Makapansgat 9.5 9.9 10 9.6 East Africa 11.5 10.4 10.5 10.6 Procercocebus antiquus (Haughton, 1925) Taung 10.9 10.5 12.1 11.2 10.4 11.4 10.4 10.4 Papio angusticeps (Broom, 1940) Kromdaai A 11.7 11.1 10.5 10.6 Cooper's 10.9 12.4 10.9 12 Sts 256 Sts 268 Sts 270 Sts 271 Sts 299 Sts 309 Sts 335 Sts 353 Sts 360 Sts 371 Sts 374 Sts 400 Sts 445 Sts 542 Sts 562 Sts 562 M2968 M2978 M2979 M2994 M3024 10.7 8.9 11 8.6 10.5 9.7 10.4 9.2 10.3 8.9 10.2 9.1 10.8 10 11.4 8.8 10.7 9.9 10.9 9.8 9.6 9.3 10.4 8.6 10.4 9.5 Makapansgat 9.5 7.6 10.8 9 East Africa 10.4 9.2 10 9.2 Procercocebus antiquus (Haughton, 1925) Taung 12.1 9.6 Papio angusticeps (Broom, 1940) Kromdaai A 12.4 9.3 BF 56787 Sts 256 Sts 261 Sts 271 Sts 309 Sts 335 Sts 369 A-B Sts 369A Sts 371 Sts 374 Sts 374 Sts 400 Sts 414A Sts 445 Sts 542 Sts 562 13 10 12.5 10.2 13.7 8.5 12.8 9.6 12.4 9.3 12.6 9.2 12.4 9 12.2 9.1 13.3 9.3 13.2 8.6 Makapansgat 11.1 9.2 11.3 7.9 13.2 8.3 12.7 9.3 East Africa 13.6 9.7 14.1 9.5 6.8 7.5 Sts 255 7 6.4 Sts 270 6.8 6.4 Sts 280 6.7 5.9 Sts 323 7 6.2 Sts 323 6.8 6.8 Sts 335 6.6 6.2 Sts 337 6.4 7.5 Sts 338 6.4 6.3 Sts 353 6.3 5.9 Sts 363 6.3 6.2 Sts 374 6.4 6.3 Sts 414A 7.3 6.1 Sts 426 6.2 6.9 Sts 445 6 7 Sts 542 7 6.2 Sts 562 7.5 7.3 1238/13 M 210 M 210 M213 Sts 342 14.1 9.6 13.9 8.2 M213 East Africa 6.4 6.5 M210 6.3 6 M210 Procercocebus antiquus (Haughton, 1925) Taung 14.6 10 Procercocebus antiquus (Haughton, 1925) Taung 7.5 7.1 Sts 352 Papio angusticeps (Broom, 1940) Sts 359 Papio angusticeps (Broom, 1940) 11.7 12.4 12.1 12.8 11.7 10.6 10.3 11.5 10.4 11 12 11.3 Sts 352 Sts 359 Sts 370A Sts 414A Sts 411A-B 14.6 10.5 M3062 Sts 563 Cooper's 12.4 9.3 11.9 10.1 12.6 9.5 12.1 10 11.6 9.7 Sts 352 Sts 352 Sts 533 Sts 359 M3062 Kromdaai A 14.3 9.9 14.6 10.7 Cooper's 14.2 10.4 Sts 370A Kromdaai A 8.3 6.4 Sts 389A 7.7 7.4 Sts 411A-B 15 10.6 Sts 563 Cooper's 7.9 6.5 14.9 9.9 Sts 563 8.1 7.2	Specimen Sts 372A Sts 372A Sts 547 Sts 547 Sts 565 Sts 565 Sts 456 A.L. 363-1b A.L. 363-15a TP9 T.17 T.10 KA 156 KA 161 KA 174 KA 194 KA 157 KA 167 CO 101 CO 135 CO 100 CO 117 Specimen MP 76 MP 76 MP 151 Sts 250 Sts 287 Sts 372A Sts 565 M2976 T.14 A.L. 363-1a A.L. 363-15a TP9 T.10 T.13 T.16 KA 161 KA 167 KA 174 KA 194 CO 101 CO 117 CO135 Specimen Sts 250 Specimen Sts 250 Specimen Sts 258 Specimen Sts 421 Specimen Sts 258 CO 109A CO135 CO101 Sts 287 Sts 322 Sts 372A Sts 372A Sts 547 Sts 547 Sts 565 M3017 A.L. 363-15a TP9 SAM 5356 T.20 T.25 T.10 T.13 T.16 KA 156 KA 161 KA 194 KA 157 CO 100 CO 117 CO105ab Sts 322 Sts 322 Sts 367 Sts 367 Sts 372A Sts 547 Sts 565 M3009 M3013 A.L. 363-1a A.L. 363-15a TP9 SAM5356 T.17 T.25 KA 156 KA 194 CO 100 Sts 270 Sts 329 Sts 331 Sts 355 Sts 355 Sts 374 A-B Sts 379 B Sts 381A Sts 390A Sts 390A Sts 409 Sts 421 Sts 469 M215 M2992 A.L. 363-1b A.L. 363-15b T.18 KA 165 Sts 258 Sts 258 Sts 329 Sts 331 Sts 374 A-B Sts 390A Sts 390A Sts 409 Sts 542 1238/12 M2984 M3019 M3021 M3030 A.L. 363-15b A.L. 363-1b Sts 269 Sts 270 Sts 280 Sts 307 Sts 313 Sts 317 Sts 329 Sts 331 Sts 348 Sts 355 Sts 355 Sts 374 A-B Sts 381A Sts 390A Sts 390A Sts 411A Sts 542 A.L. 217-8 A.L. 465-1 A.L. 363-1b A.L. 363-15b T.21 T.18 CO 101 CO 102 CO 105ab CO 109A CO 127 CO 128 CO 135 CO 115 KA 165 CO 103 CO 112 CO 115 CO 136 A-C KA 165 KA 166 A-B KA 166 A-B KA 168 CO 103 CO 135 B CO 108A CO 112 CO 136 A-C

Acknowledgements

We thank the following individuals and institutions for assistance during this project: the landowner, C. Klinkert, for permission to conduct fieldwork on her property, the South African Heritage Resource Agency (SAHRA) for their continued support in providing the excavation permit, the curators of collections: at the Ditsong: National Museum of Natural History at Pretoria in South Africa (S. Potze and M. Tawane for access to Sterkfontein, Swartkrans, Kromdraai and Taung fossil collections housed at the Plio-Pleistocene Palaeontology section; H. Fourie for access to the collection of extant primates in the Large Mammal Department), at the University of the Witwatersrand in Johannesburg, South Africa (B. Zipfel for access to the Makapan fossil collections) and at the Afrika Museum at Tervuren in Belgium (E. Gilissen and W. Vendelen in the Zoology Department for access to the collection of extant primates) and A. Lethiers (CR2P) for the help with the map of BP and one figure. We also thank the two anonymous reviewers for their comments and suggestions. The French Embassy in South Africa, the Ditsong: National Museum of Natural History, the IRL HOMEN (Hominids and Environments: Evolution of Plio-Pleistocene Biodiversity [Cradle of Humankind, South Africa]) of CNRS & NRF and the Pôle Sciences humaines et sociales, Archéologie et Patrimoine of the sous-direction de l'Enseignement supérieur et de la Recherche, French Ministry of Europe and Foreign Affairs via the Mission Paléoanthropologique Franco-Sud-Africaine (MPFSA), funded fieldwork and/or preparation and laboratory studies.