Which predators are responsible for faunal accumulations at the Late Pleistocene layers of El Harhoura 2 Cave (Témara,Morocco)?

El Harhoura 2 cave (Témara, Morocco) has yielded Aterian and Iberomaurusian lithic artifacts associated with faunal remains. Both humans and carnivores occupied this cave and non-human predator modifications occurred mainly at the end of the Late Pleistocene. Diverse faunal taxa have been identified, with a predominance of gazelles and various carnivores, particularly canids. The location of the cave and of the excavation area, at the bottom of a cliff of low elevation and in the entrance of the cave, does not correspond to a protected area for large raptor nests or a natural trap. Considering the consumed species, the type of carnivore remains, the skeletal representation of prey, taphonomic alterations such as tooth marks, semi-digested bones and destruction sequences, large canids would be the main cause for faunal modifications. However, North African fossil data attributed to hyena activities present similar results. This paper highlights the difficulty of discriminating between potential accumulators/consumers due to a lack of taphonomic reference data. This study thus demonstrates the necessity of compiling fossil records and neotaphonomic reference data for North African medium-large predators in order to better understand the taphonomic history of North African archaeological and paleontological sites.     

What happens to dead birds left in a cave and its vicinity for a quarter of a century? A taphonomic experiment

Abstract

This paper describes a bird carcass-monitoring experiment carried out in a limestone cave and its immediate vicinity in southern Poland for almost a quarter of a century. Some specimens deposited outside the cave were preserved almost intact, others were only a little weathered. Soft tissues and remnants of feathers were preserved in some of the samples inside the cave. Stages of bone modifications that can occur in a cave have been distinguished and illustrated for the first time. Also, bird bones from owl pellets have been used in a carcass-monitoring experiment for the first time. Bone modifications in the form of wrinkling, bending and/or collapsing of the bone surface were recorded only on the bones of birds that decomposed inside the cave, and not on bones that had been isolated from owl pellets prior to their deposition in the cave. Therefore, it is postulated that avian – and possibly also small mammal – bones that have been defleshed, either by people or animals, have a greater chance to survive and fossilize than bones deposited as complete carcasses in a cave. This agrees with the generally accepted notion that larger assemblages of small vertebrates in caves are usually the result of predator accumulation.     

Microvertebrates preserved in mammal burrows from the Holocene of the Argentine Pampas: a taphonomic and paleoecological approach

Microvertebrates are a major component of many assemblages recovered from the Quaternary of the Argentine Pampas. The main goal of this paper is to analyse the taphonomic history of a Holocene microfossil bonebed, recovered from the infilling of a burrow. Evidences suggest the plains vizcacha Lagostomus maximus as the putative producer of the burrow. The assemblage includes individuals belonging to different taxa of mammals (marsupials and rodents) and reptiles (snakes). Taphonomic features suggest that the accumulation inside the burrow was related to flooding processes in the plain. The burrow was a natural trap that favoured the accumulation and preservation of remains corresponding to individuals from different sources. According to the taphonomic evidence, some individuals (Lagostomus maximus, Lestodelphys halli and Serpentes indet.) died inside the burrow, whereas others (Microcavia australis, Reithrodon auritus and Ctenomys sp.) died outside the burrow, and after a time of being exposed on the surface their remains were transported by surface run-offs into the burrow. The record of Lestodelphys halli and Serpentes indet. in the burrow produced by Lagostomus maximus could be related to a circumstantial use. Mammal burrows are a significant taphonomic mode for the late Cenozoic of the Argentine Pampas.     

Late Pleistocene hyena skeleton remains of a communal/prey depot cave den in the Bohemian Mountains (Czech Republic) – its osteology,taphonomy and palaeoecology

A total of 366 bones are arranged to six incomplete individual skeletons (=MNI) of different age classes and taphonomic preservation from the Late Pleistocene Crocuta crocuta spelaea (Goldfuss 1823) hyena den Srbsko Chlum–Komín Cave, Czech Republic, Bohemian Mountains (Central Europe). The hyena assemblage counts no siblings, 37% cub, 33% early adult/subadult and 30% adult/senile individuals. A senile scavenged individual demonstrates the cannibalism within the cave. About 10% of the 3569 megafauna bones are from hyenas which larger amounts are typical at communal/prey depot den sites. This site is different to three other larger natal/birth and communal or prey depot hyena den cave sites in Czech Republic. The Chlum–Komín Cave hyena prey bone assemblage (=NISP) consists, similar as at two other Czech Republic sites, mainly of Equus ferus przewalskii (50%), including pregnant females being hunted in spring/early summer. The fewer amounts of Coelodonta antiquitatis (4%), Bison priscus (2%), Rangifer tarandus (15%), and alpine Rupicapra rupicapra (2%) or Capra ibex (1%) and the absence of mammoth correlate to hyena den bone assemblages in middle high elevated mountain regions of northern/central Europe. The rareness of cave bears in the Bohemian Mountains, on which hyenas specialized to feed additionally in European mountain regions, explains predominant predation on horses.     

Taphonomic signature of golden eagles (Aquila chrysaetos) on bone prey remains

Abstract

The golden eagle (Aquila chrysaetos) is one of the most important birds of prey in the Northern Hemisphere. This raptor is used to building large nests in high cliffs to which they return for several breeding years accumulating important amounts of their prey skeletal remains. This makes the golden eagle one of the major predators able to accumulate faunal remains in archaeological sites. Despite this fact, the taphonomic signature of golden eagles has not been properly characterized. Here we present the analysis of ingested and non-ingested faunal remains predated and accumulated by this raptor in two different nesting areas from the Iberian Peninsula. Results show how the faunal taxonomic record may vary depending on the ecological zone. Leporids and terrestrial carnivores are the best represented. The observed anatomical representation, breakage and bone surface modification patterns are discussed for different taxa. The taphonomic pattern varies depending on the type of prey and the origin of skeletal materials (non-ingested vs. pellets). Finally, after comparing our results with marks left by other predators, several characteristic features are noted to recognise golden eagles as agents of animal bones accumulations in the fossil record.     

Avian remains from the Upper Pleistocene (MIS3) site of Aguilón P-7, south of the Ebro River,Spain

Aves are represented by abundant fossil remains in Quaternary sites. Birds are well adapted to the environment they inhabit, so they make very good paleoenvironmental indicators for Quaternary sites. Here we analyse the avian remains from the Late Pleistocene (probably MIS3) site of Aguilón P-7 (AGP-7). The Pleistocene sediments fill up a shallow cave, which is located in the Zaragozan part of the Iberian Range, 55 km south of the city of Zaragoza. We have for the first time provided a taxonomic and taphonomic study of the avian assemblage of AGP-7, as well as a preliminary paleoenvironmental analysis based on these data. Nine avian taxa have been identified: Galliformes indet., Lagopus sp., Aquila chrysaetos, Gyps fulvus, Passeridae indet., Anthus sp., Prunella modularis,Sturnus cf. unicolor and Corvus monedula. The taphonomic analysis did not provide conclusive information. However, it suggests an accumulation of uneaten food remains by diurnal birds of prey. The identified taxa currently inhabit the Iberian Peninsula, populating woodland environments with rocky areas. They are found in areas with an oceanic climate, in contrast to the Mediterranean climate that now prevails in Aguilón.     

Integrated Paleoenvironmental Reconstruction and Taphonomy of a Unique Upper Cretaceous Vertebrate-Bearing Locality (Velaux,Southeastern France)

The Velaux-La Bastide Neuve fossil-bearing site (Bouches-du-Rhône, France) has yielded a diverse vertebrate assemblage dominated by dinosaurs, including the titanosaur Atsinganosaurus velauciensis. We here provide a complete inventory of vertebrate fossils collected during two large-scale field campaigns. Numerous crocodilian teeth occur together with complete skulls. Pterosaur, hybodont shark and fish elements are also represented but uncommon. Magnetostratigraphic analyses associated with biostratigraphic data from dinosaur eggshell and charophytes suggest a Late Campanian age for the locality. Lithologic and taphonomic studies, associated with microfacies and palynofacies analyses, indicate a fluvial setting of moderate energy with broad floodplain. Palynomorphs are quite rare; only three taxa of pollen grains occur: a bisaccate taxon, a second form probably belonging to the Normapolles complex, and another tricolporate taxon. Despite the good state of preservation, these taxa are generally difficult to identify, since they are scarce and have a very minute size. Most of the vertebrate remains are well preserved and suggest transport of the carcasses over short distances before accumulation in channel and overbank facies, together with reworked Aptian grains of glauconite, followed by a rapid burial. The bones accumulated in three thin layers that differ by their depositional modes and their taphonomic histories. Numerous calcareous and iron oxides-rich paleosols developed on the floodplain, suggesting an alternating dry and humid climate in the region during the Late Campanian.     

Taphonomic Analysis of the Faunal Assemblage Associated with the Hominins (Australopithecus sediba) from the Early Pleistocene Cave Deposits of Malapa,South Africa

Here we present the results of a taphonomic study of the faunal assemblage associated with the hominin fossils (Australopithecus sediba) from the Malapa site. Results include estimation of body part representation, mortality profiles, type of fragmentation, identification of breakage patterns, and microscopic analysis of bone surfaces. The diversity of the faunal spectrum, presence of animals with climbing proclivities, abundance of complete and/or articulated specimens, occurrence of antimeric sets of elements, and lack of carnivore-modified bones, indicate that animals accumulated via a natural death trap leading to an area of the cave system with no access to mammalian scavengers. The co-occurrence of well preserved fossils, carnivore coprolites, deciduous teeth of brown hyaena, and some highly fragmented and poorly preserved remains supports the hypothesis of a mixing of sediments coming from distinct chambers, which collected at the bottom of the cave system through the action of periodic water flow. This combination of taphonomic features explains the remarkable state of preservation of the hominin fossils as well as some of the associated faunal material.     

Upper Pleistocene Panthera leo spelaea (Goldfuss, 1810) remains from the Bilstein Caves (Sauerland Karst) and contribution to the steppe lion taphonomy,palaeobiology and sexual dimorphism

Remains of the steppe lion Panthera leo spelaea (Goldfuss) from historical digs in the Bilstein Caves of Warstein (Sauerland, NW Germany) are described. Their age seems to be from the Early Weichselian periods (Upper Pleistocene). Whereas the Bilstein cave was inhabited by cave bears at that time only a few hyena prey remains, were most likely imported into the cave entrance by hyenas. Bite and crush marks on a few bones of Bison priscus, Bos primigenius, Cervus elaphus, a rhinoceros Coelodonta antiquitatis vertebra and even several chewed cave bear bones prove the hyena presence which is similar to other caves in the Sauerland hyena den cave rich region. Additionally some larger wolves subspecies Canis lupusspelaeus bones were found, but only few Crocuta crocuta spelaea remains are present. After taphonomic comparisons to six other hyena and cave bear den caves of northern Germany, this cave can be classified as a cave bear den, which was briefly used by hyenas only for food storage or commuting or cave bear predation site in one part of the Cave. The lion material refers at least to one young adult lioness, one more adult female and two male lions; therefore, at minimum, the remains of four adult individuals are represented. The absence of juvenile lion material, in contrast to cave bear cub remains in the Bilstein Caves, proves that P. leo spelaea did not use this and all other caves in the region to raise their cubs. The bone material from the Bilstein Caves would prove the same hyena-lion antagonism conflict being recently proven for the Perick Caves, Balve Cave or Martins Cave well. Other situations in caves such as the Keppler Cave and the Bilstein Cave initially show the more complex taphonomic situation of lion remains in European caves, especially in cave bear dens, where they seem to have hunted periodically cave bears, such as it is already proven for hyenas in the Sauerland Karst and other caves of Europe.     

The living and the dead: How do taphonomic processes modify relative abundance and skeletal completeness of freshwater fish?

This study is designed to determine the extent to which taphonomic processes alter the taxonomic composition of fish remains in lacustrine sediments. We wish to explore information loss in a bone assemblage relative to the original, living community. We examined fish bone assemblages from lacustrine sediments along the southern shore of Lake Kinneret (Sea of Galilee) and compared them to modern living communities. For this purpose we randomly selected 24 squares, each 0.5 m² in size, and excavated them to a depth of 30–50 cm. Three lithofacies were recovered, spanning the past 1500 years (unccorected for reservoir age). The fish remains include 5037 bones and 758 scales, of which 1566 bones were identified to taxonomic group. The list of identified species was compared with the list of indigenous species known to live in Lake Kinneret in general and in a similar sandy habitat in particular. The proportion of skeletal elements found was compared with the proportion known in a complete fish. Our study indicates that differences exist between the three lithofacies in species diversity and composition, skeletal element richness, completeness, and relative abundance. In addition, the bones exhibit a clumped distribution pattern, regardless of depositional depth. From a taphonomic and paleoecological perspective, our findings demonstrate that fish remains retrieved from lacustrine sediments do not represent the composition and diversity of species as in the recent fish community.     

Taphonomic study of leporid remains accumulated by the Spanish Imperial Eagle (Aquila adalberti)

Distinguishing leporid bones accumulated by different agents such as diurnal raptors, owls, mammals and humans is essential to gain an understanding of not only human subsistence activities but also past ecology. This is particularly relevant in Iberian Palaeolithic sites where leporid remains usually constitute the most abundant taxon. Among diurnal raptors the Spanish Imperial eagle (Aquila adalberti) has been one of the most important leporid predators throughout the Iberian Peninsula. In order to investigate the taphonomic signature of this raptor, rabbit remains from 79 pellets were examined. Results show a high proportion of distal elements of the limb bones and skull. Compared with other diurnal birds of prey, the assemblages produced by this species appear to show a higher degree of breakage and corrosion from digestion. These results place this predator within a category similar to the small mammal carnivores (category 5) in terms of skeletal element abundance, breakage and digestion. It is hoped that these data will enable analysts to identify leporid fossils accumulated by the Spanish Imperial Eagle in archaeological assemblages.     

Late Pleistocene leopards as a bone accumulator: taphonomic results from S’Espasa cave and other Iberian key sites

Abstract

The Pleistocene faunal accumulations documented in caves have commonly been attributed to the activity of humans or carnivores. According to the palaeontological and archaeological literature, cave hyena (Crocuta spelaea) was the main known bone accumulator in karstic environments. However, in recent times, the role of leopards as bone accumulators has been revealed, and recent research has identified this behaviour in the Iberian Pleistocene. Moreover, there are other caves where leopard could have been claimed as an accumulator such as S’Espasa. In this work we present its taphonomic study. This cave was compared with the actualistic studies of leopards. Besides, the site of S’Espasa was compared with the other leopard dens in the Iberian Peninsula. These sites present faunal assemblages composed mainly by leopard (Panthera pardus) and Iberian wild goat (Capra pyrenaica), the bones of this ungulate present a similar pattern of bone modification by carnivores, skeletal survival rate, and bone breakage. These features indicate that goats could have been accumulated by leopards. With the data from this work and the previous ones, we try to establish a pattern that will help in the future to identify other accumulations created by this big cat.     

Taphonomy and palaeopathology in archaeozoology

Taphonomy studies post mortem damage to animal bones. Palaeopathology is the study of in vivo lesions in the skeleton. Archaeozoology uses animal remains from archaeological sites to study the relationship between people and animals. The connection between these concepts is that while all archaeozoological finds are subject to some kind of taphonomic process, only a certain number originate from diseased animals. Palaeopathology developed special features in archaeozoology, since contemporary human interference must always be reckoned with in archaeological assemblages. Animal remains occur as scattered-food refuse so that most diagnoses are isolated from their biological context. This paper reviews through various examples why a taphonomic approach to palaeopathological studies is of utmost significance.     

Modern hunting behavior in the early Middle Paleolithic: faunal remains from Misliya Cave, Mount Carmel, Israel

Understanding the behavioral adaptations and subsistence strategies of Middle Paleolithic humans is critical in the debate over the evolution and manifestations of modern human behavior. The study of faunal remains plays a central role in this context. Until now, the majority of Levantine archaeofaunal evidence was derived from late Middle Paleolithic sites. The discovery of faunal remains from Misliya Cave, Mount Carmel, Israel (>200 ka), allowed for detailed taphonomic and zooarchaeological analyses of these early Middle Paleolithic remains. The Misliya Cave faunal assemblage is overwhelmingly dominated by ungulate taxa. The most common prey species is the Mesopotamian fallow deer (Dama mesopotamica), followed closely by the mountain gazelle (Gazella gazella). Some aurochs (Bos primigenius) remains are also present. Small-game species are rare. The fallow deer mortality pattern is dominated by prime-aged individuals. A multivariate taphonomic analysis demonstrates (1) that the assemblage was created solely by humans occupying the cave and was primarily modified by their food-processing activities; and (2) that gazelle carcasses were transported complete to the site, while fallow deer carcasses underwent some field butchery. The new zooarchaeological data from Misliya Cave, particularly the abundance of meat-bearing limb bones displaying filleting cut marks and the acquisition of prime-age prey, demonstrate that early Middle Paleolithic people possessed developed hunting capabilities. Thus, modern large-game hunting, carcass transport, and meat-processing behaviors were already established in the Levant in the early Middle Paleolithic, more than 200 ka ago.     

Estimating diaphyseal length from fragmentary subadult skeletal remains: Implications for palaeodemographic reconstructions of a southern Ontario ossuary

Fragmentary skeletal remains are a significant problem for osteologists attempting to reconstruct individuals or populations. This problem is further aggravated by sites yielding commingled remains, such as are recovered from the large protohistoric and historic ossuaries from southern Ontario, for which individual methods of age estimation and sex determination cannot be used concurrently. While some attention has been given to the estimation of long bone length from fragmentary, adult remains, little attention has been given to the equally important problem of fragmentary long bones in subadult assemblages. Analysis of data on diaphyseal length is a crucial aspect of reconstructing subadult palaeodemographic profiles, particularly for ossuary collections where dental remains are not associated with individuals and are often less represented than long bones. Such analysis also aids in the assessment of conditions of past population health. This study reports the results of several regression techniques used to estimate diaphyseal length from shaft-end breadths. Data collected from two southern Ontario ossuary samples were compiled to calculate the regression equations. Reliability of these equations and implications for palaeodemographic profiles are discussed. © 1996 Wiley-Liss, Inc.     

Live and dead benthic foraminiferal assemblages from coastal environments of the Aegean Sea (Greece): Distribution and diversity

Benthic foraminiferal composition assemblages and their temporal changes, ecological indices and foraminiferal densities are used to compare three coastal environments with different physicogeographical features in the Aegean Sea (coastal environment of Avdira–Vistonikos Gulf and Kitros–Thermaikos Gulf and open lagoonal environment of Vravron–South Evoikos Gulf). Three main foraminiferal assemblages have been recognized: a) “Assemblage A”; high degree of similarity between living and dead foraminiferal species, dominated by Ammonia beccarii, Elphidium spp. and relatively abundant and diverse miliolids, b) “Assemblage B1”; intermediate degree of similarity between live and dead assemblages, characterized by highly-abundant and well-diversified foraminiferal assemblages including the algal symbiont bearing Peneroplis pertusus together with Ammonia tepida and several small epiphytic rotaliids and miliolids, and c) “Assemblage B2”; absence of living individuals, strongly dominated by the opportunistic species A. tepida. Our results suggest a good comparison between living and dead assemblages from different coastal environments in the Aegean Sea, however the prevailing environmental conditions (vegetation cover, hydrodynamics, fresh water influx) have a strong impact on the taphonomic processes.     

甘肃灵台上新世哺乳动物化石埋藏学

灵台任家坡动物群年龄组成以老年、幼年个体为主,显示动物正常死亡条件下的骨骼富集积累过程。由于动物死亡与掩埋之间时限的长短不同导致个体间风化痕迹、关节分解、破损程度不同。化石具有明显的定向性及分选性,经过季节性水流短距离搬运和较弱的分选作用。化石的埋藏特点与泛滥平原骨骼富集有较多的相似性,但沉积物组成差别很大,暗示中国红粘土物源的特殊性。     

The context of Stw 573, an early hominid skull and skeleton from Sterkfontein Member 2: taphonomy and paleoenvironment

The reconstructed taphonomic and paleoenvironmental contexts of a ca. 4 million-year-old partial hominid skeleton (Stw 573) from Sterkfontein Member 2 are described through presentation of the results of our analyses of the mammalian faunal assemblage associated stratigraphically with the hominid. The assemblage is dominated by cercopithecoids (Parapapio and Papio) and felids (Panthera pardus, P. leo, Felis caracal, and Felidae indet.), based on number of identified specimens, minimum number of elements and, minimum number of individuals. In addition, the assemblage is characterized by a number of partial skeletons and/or antimeric sets of bones across all taxonomic groups. There is scant indication of carnivore chewing in the assemblage. These observations, in addition to other taphonomic data, suggest that the remains of many animals recovered in Member 2 are from individuals that entered the cave on their own-whether accidentally by falling through avens connecting the cave to the ground surface above or by intentional entry-and were then unable to escape, rather than primarily through systematic collection by a biotic, bone-accumulating agent. The taphonomic conclusion that animals with climbing proclivities (i.e., primates and carnivores) are preferentially preserved over other taxa, ultimately because of those proclivities, urges caution in assessing the fidelity of the assemblage for reconstruction of the Member 2 paleoenvironment. With that caveat, we infer that the Member 2 paleoenvironment was typified by rolling, rock-littered and brush- and scrub-covered hills, indicated by the abundant F. caracal and cercopithecoid fossils recovered and the identified presence of the extinct Caprinae Makapania broomi. In addition, the valley bottom may have retained standing water year-round, perhaps supporting some tree cover--a setting suitable for the well-represented ambush predator P. pardus and suggested by the presence of Alcelaphini. Finally, the reconstructed taphonomic and paleoenvironmental settings of Sterkfontein Member 2 are compared to penecontemporaneous sites in South and East Africa.     

Taphonomy and palaeoecology of the late Pleistocene to middle Holocene small mammal succession of El Harhoura 2 cave (Rabat-Témara, Morocco)

The relationship between local and global climatic variations and the origin and dispersal of Homo sapiens in Africa is complex, and North Africa may have played a major role in these events. In Morocco, very few studies are specifically dedicated to small fossil vertebrates, and neither taphonomic nor palaeoecological studies have been undertaken on these taxa, particularly in archaeological contexts. The late Pleistocene to middle Holocene succession of El Harhoura 2 cave, situated in the region of Témara, yields an exceptionally rich small vertebrate assemblage. We present the results of a first systematic, taphonomic, and palaeoecological study of the small mammals from Levels 1 to 8 of El Harhoura 2. The absence of bone sorting and polishing, as well as the presence of significant traces of digestion indicate that the small mammal bones were accumulated in the cave by predators and that no water transport occurred. Other traces observed on the surface of bones consist mainly of root marks and black traces (micro-organisms or more probably manganese) which affected the majority of the material. The percentage of fragmentation is very high in all stratigraphic levels, and the post-depositional breakage (geologic and anthropogenic phenomena) obscure the original breakage patterns of bones by predators. According to the ecology of the different species present in the levels of El Harhoura 2, and by taking into account possible biases highlighted by the taphonomic analysis, we reconstruct the palaeoenvironmental evolution in the region. For quantitative reconstructions we used two indices: the Taxonomic Habitat Index (THI) and the Gerbillinae/Murinae ratio. Late Pleistocene accumulations were characterised by a succession of humid (Levels 3, 4a, 6, and 8) and arid (Levels 2?, 5, and 7) periods, with more or less open landscapes, ending in an ultimate humid and wooded period during the middle Holocene (Level 1). We discuss particular limits of our results and interpretations, due to an important lack of taxonomic, ecological, and taphonomic knowledge in North Africa.