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Mammalia, Carnivora) from Olduvai Gorge, Tanzania (Early Pleistocene

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TO L O N O G E I L C A A P I ' T A A T L E I I A Bollettino della Società Paleontologica Italiana, 55 (1), 2016, 1-9. Modena C N O A S S. P. I. New material of hyaenids (Mammalia, Carnivora) from Olduvai Gorge, Tanzania (Early Pleistocene) Marco CHERIN, Dawid A. IURINO, Jackson K. NJAU & Fidelis T. MASAO M. Cherin, Dipartimento di Fisica e Geologia, Università di Perugia, Via A. Pascoli, I-06123 Perugia, Italy; PaleoFactory, Sapienza Università di Roma, P.le A. Moro 5, I-00185 Roma, Italy; [email protected] D.A. Iurino, Dipartimento di Scienze della Terra, Sapienza Università di Roma, P.le A. Moro 5, I-00185 Roma, Italy; PaleoFactory, Sapienza Università di Roma, P.le A. Moro 5, I-00185 Roma, Italy; [email protected] J.K. Njau, Department of Geological Sciences, Indiana University, 1001 East 10th Street, IN 47405-1405 Bloomington, USA; The Stone Age Institute, Indiana, USA; [email protected] F.T. Masao, Department of History & Archaeology, University of Dar es Salaam, P.O. Box 35050, Dar es Salaam, Tanzania; [email protected] KEY WORDS - Hyaenidae, Hyaena, Crocuta, Early Pleistocene, Olduvai, Tanzania. ABSTRACT - The Hyaenidae are today a small carnivoran family including only four monospecific genera, although their paleodiversity was very high from the Miocene to the Pleistocene, with a paleobiogeographic distribution spanning Eurasia, Africa and North America. The living species Crocuta crocuta, Parahyaena brunnea and Proteles cristatus are limited to the African Continent, while the distribution of Hyaena hyaena includes northern, central and eastern Africa and southwestern Asia. The paleontological record suggests that forms very similar to the extant ones lived in Africa at least from the late Pliocene. Here we report new hyaenid craniodental material from the renowned site of Olduvai Gorge, in northern Tanzania. The fossils were found in three different Olduvai localities (FC West, MCK East and Loc. 64 in Naisiusiu area), within layers spanning stratigraphically from Bed I to Upper Bed II (Early Pleistocene). We refer the remains to Hyaena sp., Crocuta cf. ultra and Crocuta sp., supporting the occurrence of at least two hyenas (the smaller and slenderer Hyaena and the large and robust Crocuta) in the Olduvai Bed I-II carnivore guild. RIASSUNTO - [Nuovo materiale di ienidi (Mammalia, Carnivora) dalla Gola di Olduvai, Tanzania (Pleistocene Inferiore)] - Gli Hyaenidae sono oggi una piccola famiglia di carnivori comprendente solo quattro generi monospecifici, ma la loro paleodiversità è stata molto alta tra il Miocene e il Pleistocene, con una distribuzione paleobiogeografica che spaziava in Eurasia, Africa e Nord America. Le specie viventi Crocuta crocuta, Parahyaena brunnea e Proteles cristatus sono ristrette al continente africano, mentre l’areale di Hyaena hyaena comprende l’Africa settentrionale, centrale e orientale e l’Asia sudoccidentale. Il record paleontologico suggerisce che forme molto simili a quelle attuali fossero già presenti in Africa almeno dal Pliocene superiore. In questo lavoro, descriviamo nuovo materiale craniodentale di ienidi dal noto sito della Gola di Olduvai, in Tanzania settentrionale. I fossili sono stati rinvenuti in tre differenti località della Gola (FC West, MCK East e Loc. 64 nell’area di Naisiusiu), all’interno di livelli che abbracciano stratigraficamente un intervallo compreso tra il Bed I e l’Upper Bed II (Pleistocene Inferiore). I resti vengono riferiti a Hyaena sp., Crocuta cf. ultra e Crocuta sp., supportando così la presenza di almeno due iene (la più piccola e snella Hyaena e la grossa e robusta Crocuta) nell’associazione a carnivori dei Bed I-II di Olduvai. INTRODUCTION preliminary restoration and cataloguing of paleontological and archeological remains from various sites of the Gorge, Olduvai Gorge in northern Tanzania (Fig. 1) is in order to save them by human-induced and natural recognized as one of the most important paleontological destructive agents. and archeological sites in the world. The huge amount of Here we describe new hyaenid skull remains collected fossils and stone tools found in the Olduvai stratigraphic during the 2014 and 2015 field seasons from Early succession (Fig. 1) allows to reconstruct almost Pleistocene layers of three different sites of Olduvai uninterruptedly the evolution of East African ecosystems Gorge. in the last two million years, as well as some crucial phases of early human evolution. Much of the fame of Olduvai Institutional abbreviations Gorge is due to the research activity of Mary and Louis LCCO: Leakey Camp Collection Olduvai, Tanzania; Leakey, who have worked at the site for more than thirty NMDS: National Museum and House of Culture, Dar es years. Among their most important findings are Olduvai Salaam, Tanzania; UPOP: University of Perugia Olduvai Hominid (OH) 5 and OH 7, representing respectively the Project. holotypes of Paranthropus boisei (Leakey, 1959) and Homo habilis Leakey, Tobias and Napier, 1964, which were the first ancient hominin fossils discovered in East THE HYAENIDAE: FOSSIL RECORD Africa (Blumenschine et al., 2012). AND EVOLUTIONARY HISTORY Many international scientific projects are currently carried out in Olduvai Gorge, including that of the The family Hyaenidae includes only four living University of Perugia School of Paleoanthropology. This species, each assigned to its own genus: the spotted was started in 2011 as the first Italian field project at hyena Crocuta crocuta (Erxleben, 1777), the striped Olduvai and is aimed at the surface collection (Fig. 2), hyena Hyaena hyaena (Linnaeus, 1758), the brown ISSN 0375-7633 doi:10.4435/BSPI.2016.01 2 Bollettino della Società Paleontologica Italiana, 55 (1), 2016 Fig. 1 - The site of Olduvai Gorge, Tanzania. a) Stratigraphy (redrawn from Hay, 1976); b) outline map (redrawn from Hay, 1976), with location of the three localities in which the fossils described in the paper were found in 2014-2015; c) location of Olduvai Gorge within the Ngorongoro Conservation Area, northern Tanzania. Online version in color. hyena Parahyaena brunnea (Thunberg, 1820), and fact carcasses are the primary energy source for the striped the aardwolf Proteles cristatus (Sparrman, 1783). The hyena and especially the brown hyena, their feeding taxonomic status of the genus Parahyaena is discussed, habits are very opportunistic and also include many other as sometimes it is regarded as a subgenus of Hyaena food sources such as insects, fruit and small-medium (Kingdon & Hoffmann, 2013). This is mostly attributed prey (Kingdon & Hoffmann, 2013). On the other hand, to the limited knowledge on the phylogeny of hyaenids. the spotted hyena is an efficient pack hunter, capable to According to the traditional view, the genus Proteles kill large prey such as wildebeests, buffalos and zebras would be basal to the clade including Crocuta and the (Kingdon & Hoffmann, 2013). Finally, the aardwolf feeds sister taxa Hyaena and Parahyaena, thus supporting almost exclusively on termites (Holekamp & Kolowski, the inclusion of the two taxa in the same genus. On the 2009). contrary, some studies (Werdelin & Solounias, 1991) The morphology and adaptations of the four living recognize a greater affinity between the brown hyena and taxa are the remnants of the great diversity that has the spotted hyena, confirming the taxonomic validity of the characterized the Hyaenidae in the geological past. genus Parahyaena. Later molecular studies on Hyaenidae Whereas their current distribution is limited to Africa (Jenks & Werdelin, 1998; Koepfli et al., 2006) support the and - only for H. hyaena - Asia, from the Miocene to the traditional view, recognizing the basal status of Proteles Pleistocene members of this family were common in the and the sister-taxon relationship between the brown whole Eurasia and Africa, and in North America with the hyena and the striped hyena, which form a clade closely only genus Chasmaporthetes Hay, 1921 (Gilbert et al., related to Crocuta. Despite the sister-taxa relationship of 2008; Holekamp & Kolowski, 2009). The hyaenids appear Parahyaena and Hyaena, the depth of time since their split in the fossil record in the early Miocene of Europe, while (and the number of fossil taxa assigned to each genus) has the earliest occurrence in Africa dates back to the middle led some (e.g., Werdelin & Solounias, 1991) to advocate Miocene of Morocco (Werdelin & Solounias, 1991; for retaining the two genera as separate (i.e., they are Werdelin & Turner, 1996; Kingdon & Hoffmann, 2013). sister-taxa today, but only because the entire family has Werdelin & Solounias (1991) divide the Hyaenidae into so few living members). two main morphological groups: the “hyaenid” taxa (i.e., Contrary to what is commonly thought, no living taxa with morphology and ecology similar to the extant hyaenid species is an exclusive scavenger. Although in Hyaena, Parahyaena and Crocuta) and “non-hyaenid” M. Cherin et alii - Hyaenids from Olduvai Gorge 3 (approx. 10.6 Ma), but the oldest fossil species, Proteles amplidentus Werdelin & Solounias, 1991, is from the middle early Pleistocene of South Africa (Werdelin & Solounias, 1991), while some doubtful material is reported in the middle Pliocene of Laetoli, Tanzania (Werdelin & Dehghani, 2011). The hyaenid taxa more derived than Hyaenictitherium are grouped (Werdelin & Solounias, 1991) into two monophyletic clades, both characterized by a general trend of body size increasing. The first clade is formed by active hunters with adaptations to cursoriality, such as Lycyaena
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  • First Description of in Situ Primate and Faunal Remains from the Plio-Pleistocene Drimolen Makondo Palaeocave Infill, Gauteng, South Africa

    First Description of in Situ Primate and Faunal Remains from the Plio-Pleistocene Drimolen Makondo Palaeocave Infill, Gauteng, South Africa

    Palaeontologia Electronica palaeo-electronica.org 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 ABSTRACT The Drimolen palaeocave system has been actively excavated since the 1990s and has produced a demographically-diverse record of Paranthropus robustus, early Homo, and a substantial record of early Pleistocene bone tools; all recovered from the Main Quarry, a single fossil bearing deposit within the system. Early surveys identified an isolated solution-tube 55 meters west of the Main Quarry filled with decalcified matrix and fossils (the Drimolen Makondo). Recent excavations into the Makondo have started to address the geology, depositional history, and faunas of the deposits; partic- ularly whether the Makondo represents a distant uneroded part of the Main Quarry infill, or deposits in-filled into a separate entrance within the same system. We present the first description of fossil macromammalian faunas from the Makondo, excavated 2013–2014. A total of 531 specimens were recovered, 268 (50.5%) of which are taxo- nomically identifiable. The resulting list is diverse given the sample size and includes primate and carnivore taxa frequently recovered at other terminal Pliocene and earlier Pleistocene localities, as well as more rarely encountered species and elements like the first postcranial remains of the hunting hyaenid (Chasmaporthetes ?nitidula) from the Cradle. While some of the Makondo fauna overlaps with taxa recovered from the Main Quarry, there are key differences between the described samples that may reflect differences in the age of the deposits and/or taphonomic processes between these deposits at Drimolen.