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First description of in situ primate and faunal remains from the Plio-Pleistocene Drimolen Makondo palaeocave infill, Gauteng, South Africa

Douglass S. Rovinsky, Andy I.R. Herries, Colin G. Menter, and Justin W. Adams

Plain Language Abstract

This manuscript details and discusses the primary description of fossil macromammalian fauna from the new Drimolen Makondo palaeocave site, located adjacent to the hominin bearing Drimolen Main Quarry in the Cradle of Humankind World Heritage site. The faunal list is surprisingly diverse given the small sample size and scant excavation (two field seasons), includes previously unrecorded elements, and suggests a depositional time differing from that of the extensively excavated main site. As such, this manuscript touches upon and affects our knowledge of the palaeozoology, palaeobiogeography, and biochronology of the Plio-Pleistocene Cradle area.

Resumen en Español

Primera descripción de restos in situ de fauna y primates del relleno de paleocuevas Drimolen Makondo del Plio-Pleistoceno de Gauteng, Sudáfrica

El sistema de paleocuevas Drimolen ha sido activamente excavado desde la década de los 90' y se ha obtenido un registro demográficamente diverso de Paranthropus robustus, Homo primitivos y un considerable registro de herramientas de hueso de edad pleistocena temprana; todo recuperado de la Cantera Principal, un único depósito fosilífero dentro del sistema. Exámenes previos identificaron un tubo de disolución aislado a 55 m al oeste de la Cantera Principal, lleno de fósiles y matriz decalcificada (el Drimolen Makondo). Excavaciones recientes en Makondo han comenzado a centrarse en la geología, historia depositacional y faunas de los depósitos, y particularmente si Makondo representa una parte no erodada distante del relleno de la Cantera Principal, o se trata de depósitos de relleno de una entrada separada dentro del mismo sistema. Aquí presentamos la primer descripción de las faunas macromamalianas fósiles de Makondo, excavadas entre el 2013 y el 2014. Un total de 531 especímenes fueron recuperados, 268 (50,5%) de los cuales son identificables taxonómicamente. La lista resultante es diversa dado el tamaño de la muestra e incluye taxones primates y carnívoros frecuentemente recuperados en otras localidades de edad pliocena cuspidal y pleistocena temprana, así como especies menos communes y elementos tales como los primeros restos postcraneales del hiénido cazador (Chasmaporthetes ?nitidula) de la Cuna. Mientras parte de la fauna de Makondo coincide con los taxones recuperados de la Cantera Principal, existen diferencias clave entre las muestras descriptas que pueden reflejar diferencias en la edad de los depósitos y/o procesos tafonómicos entre estos depósitos en Drimolen.

Palabras clave: Cercopithecoides; Chasmaporthetes; Dinofelis; Paranthropus robustus; Metridiochoerus; Cuna de la Humanidad; Drimolen; sitios de cuevas

Traducción: Diana Elizabeth Fernández

Résumé en Français

Première description de restes fauniques (dont des primates) trouvés in situ dans le remplissage de la grotte plio-pléistocène du Makondo de Drimolen, Gauteng, Afrique du Sud

Le réseau de grottes de Drimolen a été activement fouillé depuis les années 1990 et la « Carrière principale », un dépôt fossilifère unique au sein du système, a livré un registre fossile démographiquement varié de Paranthropus robustus, Homo ancien, ainsi qu'un échantillon considérable d'outils en os datant du Pléistocène ancien. Les premières prospections ont identifié une cheminée de dissolution isolée, située 55 mètres à l'ouest de la « Carrière principale », et remplie de fossiles et de matrice décalcifiée (le Makondo de Drimolen). Des fouilles récentes dans le Makondo ont commencé à aborder les questions liées à la géologie, à l'histoire dépositionnelle, et aux faunes de ces dépôts, et en particulier la question de savoir si le Makondo représente une partie distale non érodée du remplissage de la « Carrière principale » ou des dépôts de remplissage d'une entrée séparée au sein du même réseau. Nous présentons la première description des faunes fossiles de grands mammifères du Makondo, mises à jour en 2013-2014. Au total, 531 spécimens ont été trouvés, dont 268 (50,5 %) sont taxonomiquement identifiables. La liste qui en résulte est relativement diverse étant donné la taille de l'échantillon et elle inclut des taxons de primates et de Carnivores fréquemment trouvés dans d'autres localités du Pliocène terminal et du Pléistocène ancien, ainsi que des espèces plus rares, et des éléments tels que les premiers restes postcrâniens de la hyène « chasseresse » (Chasmaporthetes ?nitidula) trouvés dans le « Berceau de l'Humanité ». Bien qu'une partie de la faune du Makondo soit commune avec celle de la « Carrière principale », des différences clés entre les échantillons décrits reflètent des différences potentielles d'âge ou de processus taphonomiques entre ces dépôts à Drimolen.

Mots-clés : Cercopithecoides ; Chasmaporthetes ; Dinofelis ; Paranthropus robustus ; Metridiochoerus ; « Berceau de l'Humanité » ; Drimolen ; sites de grottes

Translator: Antoine Souron

Deutsche Zusammenfassung

Erste Beschreibung von Primaten und Faunenüberresten in Situ aus der plio-pleistozänen Drimolen Makondo Paläohöhlen-Füllung von Gauteng, Südafrika

Das Drimolen Paläohöhlen-System wurde seit den 1990er Jahren aktiv ausgegraben und hat einen demografisch diversen Nachweis an Funden von Paranthropus robustus, frühem Homo und einen beachtlichen Satz an frühpleistozänen Knochenwerkzeugen hervorgebracht. Alle Funde wurden in der Primärhöhle gemacht, einer einzigen fossilführenden Schicht innerhalb des Systems. Während früherer Untersuchungen wurde 55m westlich der Primärhöhle eine Lösungsröhre gefunden, die mit einer dekalzifizierten Matrix und Fossilien gefüllt war (die Drimolen Makondo Füllung). Jüngste Ausgrabungen in der Makondo  befassen sich nun mit der Geologie, der Ablagerungsgeschichte und der Fauna. Insbesondere wird der Frage nachgegangen, ob die Makondo einen entfernten nicht erodierten Teil der Primärhöhlenfüllung darstellt oder ob die Ablagerungen durch einen separaten Eingang innerhalb des Systems hereinkamen. Wir präsentieren die erste Beschreibung der fossilen Makro-Säugetier-Fauna der Makando, die 2013-2014 ausgegraben wurde. Es wurden insgesamt 531 Stücke geborgen von denen 268 (50,5%) taxonomisch identifizierbar sind. Die resultierende Liste ist divers wenn man den Probenumfang bedenkt und beinhaltet Taxa von Primaten und Karnivoren, die auch häufig in anderen spätpliozänen bis frühpleistozänen Fundstellen gefunden werden. Sie beinhaltet aber auch seltenere Arten und Komponenten wie die ersten postcranialen Überreste der ausgestorben Hyäne Chasmaporthetes ?nitidula aus dem „Cradle“.Während sich einige Taxa der Makondo Fauna mit denen aus der Primärhöhle überlappen, bestehen wichtige Unterschiede zwischen den beschriebenen Proben, was Altersunterschiede der Ablagerungen und/oder Unterschiede bei den taphonomischen Prozessen zwischen diesen beiden Schichten in Drimolen wiederspiegeln könnte.

Schlüsselwörter: Cercopithecoides; Chasmaporthetes; Dinofelis; Paranthropus robustus; Metridiochoerus; Wiege der Menschheit; Drimolen; Höhlenfundstellen

Translator: Eva Gebauer

Arabic

533 arab

Translator: Ashraf M.T. Elewa

 

 

TABLE 1. Listing of fauna from the Drimolen Makondo macromammalian assemblage. NISP, number of individual specimens; MNI, minimum number of individuals.

Taxon NISP MNI
Order Primates    
  Family Cercopithecidae    
    Subfamily Colobinae    
      Cercopithecoides sp. 2 2
  Cercopithecidae indet. 13 1
           
Order Carnivora    
  Family Canidae    
    Vulpes chama 14 1
  Family Felidae    
    Dinofelis sp. 3 1
  Family Hyaenidae    
    Chasmaporthetes ?nitidula 7 1
           
Order Artiodactyla    
  Family Bovidae    
    Tribe Alcelaphini    
      Alcelaphin indet. 2 1
    Tribe Antilopini    
      Antilopin indet. 1 1
    Tribe Hippotragini    
      Hippotragus sp. 9 1
  Bovidae indet. 133 1
  Family Suidae    
    Metridiochoerus sp. 1 1
           
Order Perissodactyla    
  Family Equidae    
    cf. Eurygnathohippus cornelianus  1  1
Total       186 12

TABLE 2. Linear measurements (in mm) of Cercopithecoides sp. teeth. LL = labiolingual, BL = buccolingual, MD = mesiodistal. Boldface indicates article specimens.

Specimen Side I1 LL I1 MD I2 LL I2 MD C1 LL C1 MD C1 LL C1 MD P3 BL P3 MD P4 BL P4 MD
DNM 1 Left             5.61 4.55 5.25 7.66 6.75 7.08
DN 861 Right                 5.19   6.85 7.47
DN 889 Right                     5.49 7.57
DNM 95 Right 5.54 7.66 5.24 6.34 6.79 7.99            

TABLE 3. Measurements (in mm) of the DNM Vulpes chama dentition and other small-bodied modern and fossil canid mandibular teeth. µ, mean; σ, standard deviation. Measurement definitions after Hartstone-Rose et al., 2013. Cf. Materials and Methods for measurement abbreviations. Boldface indicates article specimens.

  Vulpes chama Vulpes chama Vulpes skinneri Canidae indet. Vulpes chama Vulpes macrotis Vulpes pallida Vulpes velox Vulpes vulpes Vulpes zerda Vulpes pattisoni Vulpes pulcher
  DNM 471-3 DNM 471-4 U.W. 88-812a DN 981b (N = 32)a (N = 4)a (N = 3)a (N = 9)a (N = 9)a (N = 10)a (N = 1)a (N = 2)c
          µ σ µ σ µ σ µ σ µ σ µ σ µ σ µ σ
p4 L 6.53 6.71 6.68   6.76 0.59 7.59 0.42 5.62 0.50 7.27 0.64 9.28 0.98 5.11 0.34     7.90 0.50
p4 W 2.83 2.81 3.12   3.16 0.41 3.17 0.34 2.49 0.23 3.21 0.26 4.04 0.48 2.19 0.30     3.20 0.00
m1 L 10.12 10.33 10.26 12.90 10.70 0.77 10.43 0.81 8.74 0.56 11.49 0.65 14.37 1.50 7.40 0.86     13.05 0.95
m1 Tri L 6.30 6.22 6.33   7.07 0.66 7.48 0.57 5.45 0.76 8.27 0.27 9.62 1.00 5.01 0.40        
m1 W 4.40 6.65 4.88 4.90 4.86 0.51 4.51 0.42 4.00 0.38 4.78 0.35 5.84 0.50 3.47 0.21     4.80 0.20
m2 L 6.36   6.47   5.79 0.61 4.54 0.25 4.91 0.26 4.92 0.51 6.34 0.42 4.71 0.99 6.96   6.55 0.55
m2 W 4.27   5.22   4.59 0.55 3.74 0.28 3.18 0.41 4.19 0.37 5.00 0.37 3.52 0.27 5.28   4.70  
Sub m1 H 14.65   14.92   16.40 1.29 13.94 1.82 12.54 1.46 15.93 1.51 20.79 1.53 10.82 0.54        
Sub m1 W 6.18   5.41   5.27 0.48 5.79 1.46 3.73 0.14 5.67 0.50 6.82 0.73 3.65 0.21        
Cor-Ang     30.50   28.03 1.41 25.66 0.79 22.79 2.70 30.01 2.08 37.89 2.25 20.53 0.94        
Con-Ang 13.63 13.82 14.42   12.70 1.07 12.01 0.93 10.57 0.51 14.33 1.51 16.69 1.11 9.47 0.44        
Con W 9.28 9.44 10.62   10.20 0.78 10.04 0.43 9.06 1.07 11.40 0.57 14.47 1.20 7.80 0.62        
Con DV 3.78 3.85 3.79   3.79 0.45 3.83 0.71 3.39 0.28 4.13 0.63 5.46 0.86 2.89 0.32        
                                         
a From Hartstone-Rose et al., 2013; b from O'Regan and Menter, 2009; c Ewer, 1956.                

TABLE 4. Measurements (in mm) of the Drimolen Makondo Dinofelis sp. and comparative metrics of Felidae distal limb elements. Cf. Materials and Methods for measurement abbreviations. Boldface indicates article specimens.

Specimen Taxon Side Element TL PW PH SW SH DW DH
DNM 2 Dinofelis sp. Left MT II 83.2 14.2 23.1 12.0 10.6 18.0 17.4
DN 12 Dinofelis aff. piveteaui Left MT II 76.7 12.7   11.2 11.6 16.6 15.7
KB 5339 A Megantereon cultridens Left MT II 71.5 11.2 19.6 8.9 9.3 14.2 14.4
KB 5339 B Megantereon cultridens Left MT V 69.8 14.4 14.6 7.8 8.1 13.1  
KB 5539 C Megantereon cultridens Left MT IV 80.1 15.9 18.5 10.9 10.7 14.5 13.8
SK 1848 Dinofelis barlowi Right MT IV   15.1 17.6 10.2 10.6    
DN 2092 Dinofelis aff. piveteaui Left Calcaneus 75.5            
DN 2149c Dinofelis aff. piveteaui Right Calcaneus 84.0            
DNM 54 Dinofelis sp. Left Calcaneus 90.8            
UCMP 88783 a Dinofelis barlowi   Calcaneus 78.0            
UCMP 88784 a Dinofelis barlowi   Calcaneus 81.9            
KNM-ER 722Q a Dinofelis aff. piveteaui   Calcaneus 71.0            
(BPI) M 16201 a Dinofelis darti   Calcaneus 84.9            
KNM-LT 23696 a Dinofelis sp.   Calcaneus 83.4            
KNM-LT 23932 a Dinofelis sp.   Calcaneus 81.3            
Extant (N = 20) b Panthera leo   Calcaneus 103.2            
Extant (N = 14) b Panthera pardus   Calcaneus 64.3            
                     
a From Werdelin and Lewis, 2001, b from Lewis, 1995.

TABLE 5. Measurements (in mm) of the Drimolen Makondo Chasmaporthetes ?nitidula and comparative Hyaenidae distal limb elements. Cf. Materials and Methods for measurement abbreviations. Boldface indicates article specimens.

Specimen Taxon Element TL PW PH SW SH DW DH VL VW
DNM 3-4 Chasmaporthetes ?nitidula Fourth Metatarsal   9.8 19.5            
Extant a Crocuta crocuta (N = 2) Fourth Metatarsal 88.9 9.9 15.4 9.0 8.3 10.8      
Extant a Parahyaena brunnea (N = 1) Fourth Metatarsal 84.7 8.8 14.0 9.0 8.3        
Extant a Hyaena hyaena (N = 1) Fourth Metatarsal 88.4 8.3 14.2 6.2 6.3 8.7      
                       
IVPP V18567.1 b Chasmaporthetes gangriensis Second Metatarsal >80.0     9.5   13.0      
                       
DNM 3-3 Chasmaporthetes ?nitidula Proximal Phalanx 35.3 13.9 12.0 7.7 7.3 11.4 8.0    
DNM 3-5 Chasmaporthetes ?nitidula Proximal Phalanx 35.4 13.0 11.2 7.2 6.6 10.4 8.0    
TRO 1671 a Chasmaporthetes ossifragus Proximal Phalanx 37.6 17.1 18.0 12.5 9.5 14.6      
UF 27377 a Chasmaporthetes ossifragus Proximal Phalanx 38.6 12.4 13.4 7.0 8.5 11.3      
IVPP V18567.2 b Chasmaporthetes gangriensis Proximal Phalanx 34.4 12.6   7.4   10.8      
DN 2033 c Hyaenidae sp. Proximal Phalanx           11.7 8.5    
DN 562 c Hyaenidae sp. Proximal Phalanx 29.3 11.4 9.8 7.6 6.7 9.4 6.4    
SKX 28181 Hyaenidae sp. Proximal Phalanx 29.7 13.3 10.9 8.6 7.3 10.9 7.3    
SKX 29031 d Hyaenidae sp. Proximal Phalanx 29.9 13.4 12.0 9.1 8.2 11.5 8.3    
SKX 27170 d Hyaenidae sp. Proximal Phalanx 29.9 13.5 12.0 9.0 8.2 11.4 8.3    
Extant a Crocuta crocuta (N = 3) Proximal Phalanx 27.4 9.8 9.8 6.2 6.5 9.1      
Extant a Parahyaenabrunnea (N = 1) Proximal Phalanx 25.2 8.7 8.7 4.4 5.7 7.7      
Extant a Hyaenahyaena (N = 1) Proximal Phalanx 24.7 9.2 7.7 5.6 6.3 7.9      
                       
DNM 3-1 Chasmaporthetes ?nitidula Intermediate Phalanx   11.0 8.2            
DNM 3-2 Chasmaporthetes ?nitidula Intermediate Phalanx 22.4 11.9 9.0 7.5 5.3 11.6 7.6    
IVPP V18567.3 b Chasmaporthetes gangriensis Intermediate Phalanx 24.8 11.3   8.1   11.9      
SKX 36942 d Parahyaena brunnea Intermediate Phalanx 28.3 10.7 9.8 7.4 6.1   7.1    
                       
DNM 3-6 Chasmaporthetes ?nitidula Distal Phalanx 19.2 8.7 13.4     5.3   10.7 7.3
DNM 3-7 Chasmaporthetes ?nitidula Distal Phalanx 20.7 10.3 14.5     5.9   10.8 7.8
                       
a From Berta, 1981; b from Tseng et al., 2013; c from O'Regan and Menter, 2009; d from Turner, 1993.
 

FIGURE 1. 1, Location of Drimolen compared to Pretoria in the NE and Krugersdorp in the SW as well as other fossil sites in the UNESCO Cradle of Humankind World Heritage Area (Google Earth). 2, Google Earth image of the Drimolen hominin site showing the relationship of the Main Quarry (DMMQ) site from which the hominins have been recovered and the new Makondo site (DMK) that is the focus of this paper. T represents the location of Andre Keyser's preliminary excavation trench, the white arrow indicates the view in Figure 1.3. 3, the Makondo excavations in progress in 2014 showing the various Makondo features from which the fossils were recovered and looking east. In Figure 1.3 (4) denotes the location of Figure 1.4. 4, In situ fossils during excavation in 2014, showing the heavy manganese staining and concentration of fossils.

figure 1

FIGURE 2. The Drimolen Makondo deposit and features at the end of the 2014 excavation season. Fossils were recovered from decalcified deposits adhering to the walls of the Main Makondo in 2013 and mainly from within the northern and eastern portions of the Eastern Makondo in 2014. While the fossils are defined as having come from different makondo features these solution tubes have all been formed within the same depositional unit of the Drimolen Makondo.

figure 2

FIGURE 3. Cercopithecoides sp. craniodental specimens from the Drimolen Makondo. 1 and 2, DNM 1, partial mandible; 1, dorsal and 2, right lateral views. 3 and 4, DNM 95, right premaxilla and maxilla; 3,labial and 4, lingual views. Scale bars equal 1 cm.

figure 3

FIGURE 4. Vulpes chama craniodental specimens from the Drimolen Makondo. 1 and 2, DNM 471-1, partial cranium; 1, dorsal and 2, left lateral views. 3 and 4, DNM 471-2, right maxilla; 3, buccal and 4, occlusal views. 5 and 6, DNM 471-3, right mandible; 5, buccal and 6, lingual views. Scale bars equal 1 cm.

figure 4

FIGURE 5. Dinofelis sp. postcranial specimens from the Drimolen Makondo. 1-6, DNM 2, left second metatarsal; 1, proximal and 2, distal articular surfaces, 3, medial, 4, dorsal, 5, ventral, and 6, lateral views. 7, DNM 54-1 and 54-2, left calcaneus in articulation; anterior view. Scale bars equal 1 cm.

figure 5

FIGURE 6. Chasmaporthetes ? nitidula postcranial specimens from the Drimolen Makondo. 1-5, DNM 3-4, partial left fourth metatarsal; 1, proximal articular surface, 2, lateral, 3, ventral, 4, dorsal, and 5, medial views. 6-8, DNM 3-3, pedal proximal phalanx; 6, dorsal, 7, lateral, and 8, ventral views. 9-11, DNM 3-2, pedal middle phalanx; 9, dorsal, 10, lateral, and 11, ventral views. 12-14, DNM 3-6, pedal terminal phalanx; 12, dorsal, 13, lateral, and 14, ventral views. Scale bars equal 1 cm.

figure 6

FIGURE 7. Artiodactyla and Perissodactyla from the Drimolen Makondo assemblage. 1 and 2, DNM 143-1, Hippotragus sp. partial horn core; 1, lateral and 2, anterior views. 3-5, DNM 57, Metridiochoerus sp. right maxillary third premolar; 3, lingual, 4, occlusal, and 5, buccal views. 6 and 7, cf. Eurygnathohippus cornelianus left proximal fourth metatarsal; 6, medial and 7, posterior views. Scale bars equal 1 cm.

figure 7

 

author1Douglass S. Rovinsky
Biology Department
Grand Valley State University
Allendale, Michigan 49401
USA
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Douglass S. Rovinsky received his B.Sc. in biology from Grand Valley State University in 2011. His research currently focuses on the comparative anatomy and functional morphology of carnivoran postcrania. He is currently an Adjunct Instructor in the Biology Department of Grand Valley State University.

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author2Andy I.R. Herries
The Australian Archaeomagnetism Laboratory
Dept. Archaeology and History
La Trobe University, Melbourne Campus
Bundoora, Victoria 3086
Australia
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and Centre for Anthropological Research
Humanities Research Village
University of Johannesburg
Auckland Park 2006
South Africa

Dr. Andrew I.R. Herries obtained his Ph.D. at the University of Liverpool Geomagnetism Laboratory and Dept. Archaeology in 2003 working on a magnetostratigraphy of South African early hominin palaeocaves. He remains an Honorary Research Fellow of the Geomagnetism Laboratory at Liverpool through the School of Environmental Sciences. He has worked in South Africa since 1997 and has a wide ranging background in the excavation of palaeolithic and fossil bearing caves, palaeoanthropology, geoarchaeology, speleology, geochronology, karst geology, archaeological geophysics, & archaeomagentism (inc. Palaeomagnetism, Mineral [Rock] Magnetism & Environmental Magnetism). He is the head of the Australian Archaeomagnetism Laboratory (TAAL) and an Associate Professor and ARC Future Fellow in the Geochronology of Human and Marsupial Evolution in the Dept. Archaeology, Environment and Community Planning at La Trobe University.

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author3Colin G. Menter
Centre for Anthropological Research, Humanities Research Village
University of Johannesburg
Auckland Park 2006
South Africa
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Dr. Colin G. Menter received his Ph.D. from the University of the Witwatersrand and currently is director and project leader of the Drimolen hominin site through the Centre for Anthropological Research at the University of Johannesburg. His research focuses on the palaeobiology of hominins and the site-based significance of the Drimolen deposits for understanding the early Pleistocene landscape of the Cradle of Humankind UNESCO World Heritage Site.

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author4Justin W. Adams
Department of Anatomy and Developmental Biology
Monash University
Clayton, Victoria 3800
Australi
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Dr. Justin W. Adams received his Ph.D. in physical anthropology from Washington University in St. Louis in 2006. His research focus is on the evolution of African mammals in southern Africa and the paleoecology and taphonomy of the South African primate-bearing cave deposits. He has excavated and described mammalian assemblages from several Pliocene and Pleistocene fossil localities including Gondolin, Luleche, and Hoogland. He is currently a Senior Lecturer in the Department of Anatomy and Developmental Biology at Monash University.