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In South Africa, the contemporaneity of various Hominin species in the Pleistocene fossil record makes it complex to understand the evolutionary history and subsistence behaviors of these lineages. Understanding the environments in which they lived and how their remains got accumulated in the caves is required to provide reliable answers to these issues. Here we present the results of the analysis of a rich micromammal assemblage from Early Pleistocene Cooper’s D site, which has yielded remains attributed to the species Paranthropus robustus. Taphonomic analysis indicates that these remains were likely accumulated by species of the Family Tytonidae, most likely Tyto alba. The specimens have undergone post-predation modifications, showing a high degree of breakage probably resulting from trampling, as well as severe manganese infiltration which obscures the reading of bone surfaces and does not allow to detect possible weathering effects. The overrepresentation of the mandibles in the assemblage raises questions about supplementary post-depositional modifications and reflects the effects of an unknown taphonomic process. The association of micromammals supports previous interpretations of the environment at Cooper’s as a mosaic landscape dominated by grassland, also indicating a strong woody component near a permanent water source. With the addition of micromammals to the list of studied taxa, the Early Pleistocene site of Cooper’s Cave becomes one of the best documented sites from the region of the Cradle of Humankind in terms of species richness and palaeoecological inferences.

International audience; The simultaneous presence of several hominin species in South Africa during the Plio-Pleistocene, especially the first representatives of the genus Homo, Paranthropus and possibly late surviving Australopithecines, makes it difficult to understand the evolutionary history of these lineages, as well as their technical and subsistence behaviors. It then appears necessary to better characterize the occupations related to each species, and in particular the environment in which each of them has evolved. Small mammals such as rodents are good palaeoecological markers, but remain poorly studied in early hominin sites in South Africa. In order to understand the evolutionary dynamics of this time, the MiCoTaph project aims to reconstruct the taphonomic and palaeoenvironmental context of the Paranthropus robustus occupations of Cooper's D (Bloubank Valley, South Africa, < 1.4 Ma). An interdisciplinary approach is applied using taxonomic, taphonomic and palaeoecological methods to analysis the micromammal remains.

Panthera leo (Linnaeus, 1758) (Lion) MATERIAL EXAMINED. — Postcranial: CD 10783 a right 1 st Metacarpal; CD 3879 and CD 10814 both 1 st phalanges. DIAGNOSIS AND DISCUSSION CD 10783 is a very large 1 st metacarpal. Illustrations of Dinofelis piveteaui (Werdelin & Lewis 2001: fig. 19G) and Homotherium (Ballesio 1963) show that the facet for the 2nd metacarpal is medially placed in these machairodonts while in CD 10783 and the pantherines it covers much of the proximal surface. In the three lion MC1s available for comparison, it closely matches AZ 771, except that the fossil is from a larger animal (total length = 44.3 mm, distal breadth 18.3 mm), although it differs from BPIc186 and AZ 421 (also lions). CD 3879 is slightly damaged proximally and CD 10814 is complete. They are both from large, lion-sized felids, although the depressions on the medial and lateral sides of the distal articulation are much deeper than those seen in the modern lion. They are considerably larger than the phalanges identified as Dinofelis from Kromdraai B (KB 6036, KB 6037, KB 6038) and do not appear to be Homotherium (Ballesio 1963). These specimens are all referred to Panthera leo, but note comments below regarding large pantherines in Africa., Published as part of O'Regan, Hannah J. & Steininger, Christine, 2017, Felidae from Cooper's Cave, South Africa (Mammalia: Carnivora), pp. 315-332 in Geodiversitas 39 (2) on page 325, DOI: 10.5252/g2017n2a8, http://zenodo.org/record/5206125, {"references":["LINNAEUS C. 1758. - Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. 10 th ed., Stockholm.","WERDELIN L. & LEWIS M. E. 2001. - A revision of the genus Dinofelis (Mammalia, Felidae). Zoological Journal of the Linnean Society 132: 147 - 258. https: // doi. org / 10.1111 / j. 1096 - 3642.2001. tb 02465. x","BALLESIO R. 1963. - Monographie d'un Machairodus du gisement villafranchien de Seneze: Homotherium crenatidens Fabrini. Travaux du Laboratoire de Geologie de la Faculte des Sciences de Lyon, n. s., 9: 1 - 129."]}

Genus Dinofelis Zdansky, 1924 TYPE SPECIES. — Dinofelis abeli Zdansky, 1924 (junior synonym of D. cristata) by original designation., Published as part of O'Regan, Hannah J. & Steininger, Christine, 2017, Felidae from Cooper's Cave, South Africa (Mammalia: Carnivora), pp. 315-332 in Geodiversitas 39 (2) on page 316, DOI: 10.5252/g2017n2a8, http://zenodo.org/record/5206125, {"references":["ZDANSKY O. 1924. - Jungtertiare Carnivoren Chinas. Paleontologia Sinica, series C 2: 1 - 149."]}

The Cooper’s Cave System has produced a diverse fossil assemblage including the remains of Paranthropus robustus Broom, 1938, and early Homo. The majority of the faunal remains come from Cooper’s D, which dates to ~1.5 – 1.4 Ma. Here we describe 158 craniodental and postcranial felid fossils from Cooper’s D, including Dinofelis cf. aronoki. These fossils indicate the presence of four large felid genera at Cooper’s D: Dinofelis, Megantereon, Panthera (two species) and Acinonyx, plus two smaller taxa: Caracal and Felis. This assemblage may mark the first appearance of the modern cheetah Acinonyx jubatus (Schreber, 1775) in Africa, as well the first occurrence of the East African species Dinofelis cf. aronoki in southern Africa. This taxon appears intermediate in features between Dinofelis barlowi (Broom, 1937) and Dinofelis piveteaui (Ewer, 1955). We compare the Cooper’s D felid assemblage with those from other sites in the Cradle of Humankind, Gauteng, and discuss several scenarios for the evolution of the genus Dinofelis in eastern and southern Africa.

Panthera pardus (Linnaeus, 1758) (Leopard) MATERIAL EXAMINED. — Postcranial. CD 3277, left proximal radius (Table 3); CD 7369, left radius – distal epiphysis only (Table 3); CD 1526, right ulna – distal epiphysis only; CD 1956, patella; CD 5957, 3rd Metatarsal (Table 3); CD 3836, 4th Metatarsal (Table 3); CD 8288, and CD 1537, both 1 st phalanges. DESCRIPTION AND TAXONOMIC ASSIGNMENT Forelimb CD 3277 is a proximal radius. It is smaller than the East African Dinofelis specimens illustrated in Werdelin & Lewis (2001) although the angle between the shaft and head is similar. The Kromdraai Megantereon radii (KB 5333O and KB 5336) are both larger and have a more robust radial tuberosity than is seen in CD 3277. A distal radial epiphysis (CD 7369) has a large ulnar facet, indicating that is it not a cheetah, and overall shape of the carpal facets is squarish, whereas it is more rectangular in Dinofelis and Megantereon. Morphology of the radius, both proximally and distally appears to be quite variable in modern leopards, particularly the outline and depth of the proximal articulation, but CD 3277 and CD 7369 are both most similar to modern leopards. An isolated distal epiphysis from a right ulna (CD 1526) is also assigned to P. pardus, as the styloid process in Dinofelis is much more bulbous and Dinofelis is larger overall. Hindlimb CD 1956 is a tear-drop shaped patella with some damage to the dorsal surface. It is substantially smaller than KB 5377, a patella described as cf. Megantereon from Kromdraai B. No Dinofelis patellae were available for study, but CD 1956 is very similar to a modern leopard (AZ 420) and is therefore been referred to that species. Two metatarsals have also been referred to P. pardus. CD 5957 is a complete right 3rd metatarsal, broken into three pieces. It is very gracile in comparison with Dinofelis, of a similar length but slenderer. The posterior facet of the MT4 articulation is curved, in DN17 it is not curved, while in KB 5334B (Megantereon whitei) it is flat and angled medially. Again the morphology of the leopards appears to be highly variable, but for a medium-sized felid this specimen is much more like P. pardus than any of the other similar sized species. CD 3836 is the proximal articulation plus 1/3rd of the shaft of a left 4th metatarsal. It has some slight pathological bone growth on the dorsal surface and the articulation for the 5 th metacarpal. Despite this it is clear that it is neither Dinofelis nor Megantereon – Dinofelis (DN 14) has a much more rounded articulation for the 3rd metatarsal, while this articulation is flatter and the proximal articulation is a little larger in Megantereon (KB 5339C). CD 1537 is a complete 1 st phalanx with a small chip from the dorsal surface, while CD 8288 has a small amount of damage on the distal condyle. Both are very good matches for modern leopard., Published as part of O'Regan, Hannah J. & Steininger, Christine, 2017, Felidae from Cooper's Cave, South Africa (Mammalia: Carnivora), pp. 315-332 in Geodiversitas 39 (2) on page 325, DOI: 10.5252/g2017n2a8, http://zenodo.org/record/5206125, {"references":["LINNAEUS C. 1758. - Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. 10 th ed., Stockholm.","WERDELIN L. & LEWIS M. E. 2001. - A revision of the genus Dinofelis (Mammalia, Felidae). Zoological Journal of the Linnean Society 132: 147 - 258. https: // doi. org / 10.1111 / j. 1096 - 3642.2001. tb 02465. x"]}

Megantereon whitei (Broom, 1937) (Fig. 5) MATERIAL EXAMINED. — Craniodental. CD 5963, right posterior mandible fragment with M 1 roots (Fig. 5A); CD 5997, left mandible from symphysis to M 1 (Fig. 5B, and see Hartstone-Rose et al. 2007); CD 10452, damaged right M 1 (Fig. 5C, D; Table 2); Postcranial: CD 3221, left proximal tibial epiphysis; CD 7336, left navicular; CD 5978, right navicular. DESCRIPTION AND TAXONOMIC ASSIGNMENT The three craniodental specimens are clearly attributable to Megantereon (Fig. 5 A-D). The most complete, CD 5997, is shown in Fig. 5B and fully described by Hartstone-Rose et al. (2007). The other two specimens are also from the lower jaws. CD 5963 is a posterior fragment of mandible, broken horizontally above the condyle and also anterior to the M 1 alveolus (Fig. 5A). The masseteric fossa is shallow and extends to the posterior root of the M 1, but the most notable feature is the very small distance (22 mm) between the angle of the ramus and the condylar process. The coronoid process must have been correspondingly small, and this indicates that the specimen can only have belonged to a very small machiarodont. CD 5963 is of similar size to CD5997, although the carnassial in CD 5963 may have been slightly larger. CD 10452 is an unworn M 1 that is broken across the protoconid (Fig. 5C, D). The paraconid is small (length: 8.7 mm) with a relatively larger protoconid. It is much smaller than KA64, and is most similar to the heavily damaged type specimen of Megantereon whitei (TM 856) from Schurveberg (Broom 1937; Turner 1987b). In contrast, both the P 4 and M 1 of CD 5997 are smaller than those of TM 856. As discussed by Hartstone-Rose et al. (2007) the previously known Megantereon whitei material from Coopers D is very small, and these specimens fit within that hypodigm. They most closely fit with the morphology of the type specimen of M. whitei, and there is growing consensus that M. whitei is the only Pleistocene species of the genus Megantereon in Africa (Palmqvist et al. 2007; Werdelin & Peigné 2010). Therefore the Cooper’s D specimens are assigned to this species. Three postcranial specimens have also been assigned to M. whitei. An isolated proximal epiphysis from a left tibia with some damage to the ventral edge (CD 3221) is an excellent match for KB 5333M, a partial skeleton of Megantereon whitei published by Vrba (1981). The two naviculae (CD 7336 and CD 5978) may be antimeres and are very similar to the illustrations of M. cultridens (Cuvier, 1824) from Senéze (Christiansen & Adolfssen 2007) and KB 6018 (Megantereon whitei). In comparison with DN 2571 (here referred to D. cf. aronoki) the two Cooper’s D naviculae are smaller, not so thick and have less clearly defined facets., Published as part of O'Regan, Hannah J. & Steininger, Christine, 2017, Felidae from Cooper's Cave, South Africa (Mammalia: Carnivora), pp. 315-332 in Geodiversitas 39 (2) on page 323, DOI: 10.5252/g2017n2a8, http://zenodo.org/record/5206125, {"references":["BROOM R. 1937. - On some new Pleistocene mammals from limestone caves of the Transvaal. South African Journal of Science 33: 750 - 768.","TURNER A. 1987 b. - Megantereon cultridens (Cuvier) (Mammalia, Felidae, Machairodontinae) from Plio-Pleistocene deposits in Africa and Eurasia, with comments on dispersal and the possibility of a New World Origin. Journal of Paleontology 61: 1256 - 1268. https: // doi. org / 10.1017 / S 0022336000029632","PALMQVIST P., TORREGROSA V., PEREZ- CLAROS J. A., MARTINEZ- NAVARRO B. & TURNER A. 2007. - A re-evaluation of the diversity of Megantereon (Mammalia, Carnivora, Machairodontinae) and the problem of species identification in extinct carnivores. Journal of Vertebrate Paleontology 27: 160 - 175. https: // doi. org / 10.1671 / 0272 - 4634 (2007) 27 [160: AROTDO] 2.0. CO; 2","WERDELIN L. & PEIGNE S. 2010. - Carnivora, in WERDELIN L. & SANDERS W. J. (eds), Cenozoic Mammals of Africa. University of California Press, Berkeley: 602 - 657.","VRBA E. S. 1981. - The Kromdraai australopithecine site revisited in 1980; recent investigations and results. Annals of the Transvaal Museum 33: 17 - 60.","CUVIER G. 1824. - Recherches sur les ossemens fossils, ou l'on retablit les charateres de plusieurs animaux don't les revolutions du globe ont detruit les especes. Vol. 4, 514 p.","CHRISTIANSEN P. & ADOLFSSEN J. S. 2007. - Osteology and ecology of Megantereon cultridens SE 311 (Mammalia; Felidae; Machairodontinae), a sabrecat from the Late Pliocene - Early Pleistocene of Seneze, France. Zoological Journal of the Linnean Society 151: 833 - 884. https: // doi. org / 10.1111 / j. 1096 - 3642.2007.00333. x"]}

Felis silvestris lybica Forster, 1780 (African wild cat) MATERIAL EXAMINED. — CD 691, left CS (Table 1). DESCRIPTION AND TAXONOMIC ASSIGNMENT This is a small upper canine with a broken tip. One lingual and two buccal grooves are visible in the enamel, it is clearly a small felid and based on its size, it is most likely to be F. s. lybica the African wildcat., Published as part of O'Regan, Hannah J. & Steininger, Christine, 2017, Felidae from Cooper's Cave, South Africa (Mammalia: Carnivora), pp. 315-332 in Geodiversitas 39 (2) on page 326, DOI: 10.5252/g2017n2a8, http://zenodo.org/record/5206125, {"references":["FORSTER G. 1780. - Beschreibung des rothen Baumlaufers von der Insel O-Waihi, in BUFFON G. L. L. de (ed.), Naturgeschichte der vierfussigen Thiere 6. Joachim Pauli, Berlin, 313 p."]}

Acinonyx jubatus (Schreber, 1775) MATERIAL EXAMINED. — Craniodental. CD 3871, left P4 (Fig. 5H, I; Table 1); CD 9614 left I3. DIAGNOSIS AND DISCUSSION CD 3871 is an almost complete P4, just lacking the mesial border of the protocone.Despite this it can be seen that the protocone was much reduced in comparison with the pantherines. The ectoparastyle is very large, in contrast to Megantereon where there is no ectoparastyle (Christiansen & Adolfssen 2007; KA 64 pers. obs.). Other than the protocone being slightly more anteriorly placed in CD 3871 it is a very good match for the modern cheetah. The isolated lower incisor CD 9614 has a clear accessory cusp on the buccal surface and is a robust tooth with a relatively short crown. Other than the slight difference in the protocone position on the P4, the Cooper’s D specimens match those of the modern cheetah, and are referred to this species., Published as part of O'Regan, Hannah J. & Steininger, Christine, 2017, Felidae from Cooper's Cave, South Africa (Mammalia: Carnivora), pp. 315-332 in Geodiversitas 39 (2) on page 326, DOI: 10.5252/g2017n2a8, http://zenodo.org/record/5206125, {"references":["SCHREBER J. C. D. 1775. - Die Saugethiere in Abbildungen nach der Natur, mit Beschreibungen. Vol. 2. Walther, Erlangen.","CHRISTIANSEN P. & ADOLFSSEN J. S. 2007. - Osteology and ecology of Megantereon cultridens SE 311 (Mammalia; Felidae; Machairodontinae), a sabrecat from the Late Pliocene - Early Pleistocene of Seneze, France. Zoological Journal of the Linnean Society 151: 833 - 884. https: // doi. org / 10.1111 / j. 1096 - 3642.2007.00333. x"]}

International audience; The Plio-Pleistocene karstic sedimentary deposits of Sterkfontein Cave, South Africa, yielded numerous fossil primate specimens embedded in blocks of indurated breccia, including the partial cercopithecoid cranium labelled STS 1039. Because the surrounding matrix masks most of its morphology, the specimen remains taxonomically undetermined. While the use of X-ray microtomography did not allow extracting any structural information about the specimen, we experimented a new investigative technique based on neutron microtomography. Using this innovative approach, we successfully virtually extracted, reconstructed in 3D and quantitatively assessed the preserved dentognathic structural morphology of STS 1039, including details of its postcanine maxillary dentition. Following comparative analyses with a number of Plio-Pleistocene and extant cercopithecoid taxa, we tentatively propose a taxonomic attribution to the taxon Cercopithecoides williamsi. Our experience highlights the remarkable potential of this novel imaging method to extract diagnostic information and to identify the fossil remains embedded in hard breccia from the South African hominin-bearing cave sites.

A partial hominin skull (COB 101) was identified in the fossil collections of the Transvaal Museum, Pretoria, attributed to the Cooper's Cave site in South Africa. The find represents the most complete hominin specimen recovered from localities at this site to date. COB 101 comprises the supraorbital, zygomatic, infraorbital and nasoalveolar regions of the right side, and the right upper third premolar. The specimen has undergone post-depositional distortion that resulted in the flattening of the facial structures. Here we describe and compare COB 101 with other hominin material from Africa and find that this specimen shares numerous diagnostic features with Paranthropus robustus. The discovery of COB 101 augments the number of specimens attributed to this species from other South African sites and other Cooper's Cave localities.