- •Preface
- •Contents
- •Contributors
- •1 Introduction: Azokh Cave and the Transcaucasian Corridor
- •Abstract
- •Introduction
- •History of Excavations at Azokh Caves
- •Excavations 1960–1988
- •Excavations 2002–2009
- •Field Seasons
- •2002 (23rd August–19th September)
- •2003 (4th–31st August)
- •2004 (28th July–6th August)
- •2005 (26th July–12th August)
- •2006 (30th July–23rd August)
- •2007 (9th July–4th August)
- •2008 (8th July–14th August)
- •2009 (17th July–12th August)
- •Correlating Huseinov’s Layers to Our Units
- •Chapters of This Book
- •Acknowledgments
- •References
- •Abstract
- •Introduction
- •Azokh 1
- •Sediment Sequence 1
- •Sediment Sequence 2
- •Discussion on the Stratigraphy of Azokh 1
- •Azokh 2
- •Azokh 5
- •Discussion on the Stratigraphy of Azokh 5
- •Conclusions
- •Acknowledgments
- •References
- •3 Geology and Geomorphology of Azokh Caves
- •Abstract
- •Introduction
- •Geological Background
- •Geomorphology of Azokh Cave
- •Results of the Topographic Survey
- •Azokh 1: Main Entrance Passageway
- •Azokh 2, 3 and 4: Blind Passages
- •Azokh 5: A Recently Discovered Connection to the Inner Chambers
- •Azokh 6: Vacas Passageway
- •Azokh I: The Stalagmite Gallery
- •Azokh II: The Sugar-Mound Gallery
- •Azokh III: The Apron Gallery
- •Azokh IV: The Hall Gallery
- •Results of the Geophysical Survey
- •Discussion
- •Conclusions
- •Acknowledgments
- •References
- •4 Lithic Assemblages Recovered from Azokh 1
- •Abstract
- •Introduction
- •Methods of Analysis
- •Results
- •Unit Vm: Lithic Assemblage
- •Unit III: Lithic Assemblage
- •Unit II: Lithic Assemblage
- •Post-Depositional Evidence
- •Discussion of the Lithic Assemblages
- •Comparison of Assemblages from the Earlier and Current Excavations
- •Chronology
- •Conclusions
- •Acknowledgements
- •References
- •5 Azokh Cave Hominin Remains
- •Abstract
- •Introduction
- •Hominin Mandibular Fragment from Azokh 1
- •Discussion of Early Work on the Azokh Mandible
- •New Assessment of the Azokh Mandibular Remains Based on a Replica of the Specimen
- •Discussion, Azokh Mandible
- •Neanderthal Remains from Azokh 1
- •Description of the Isolated Tooth from Azokh Cave (E52-no. 69)
- •Hominin Remains from Azokh 2
- •Human Remains from Azokh 5
- •Conclusions
- •Acknowledgements
- •References
- •6 The New Material of Large Mammals from Azokh and Comments on the Older Collections
- •Abstract
- •Introduction
- •Materials and Methods
- •General Discussion and Conclusions
- •Acknowledgements
- •References
- •7 Rodents, Lagomorphs and Insectivores from Azokh Cave
- •Abstract
- •Introduction
- •Materials and Methods
- •Results
- •Unit Vm
- •Unit Vu
- •Unit III
- •Unit II
- •Unit I
- •Discussion
- •Conclusions
- •Acknowledgments
- •8 Bats from Azokh Caves
- •Abstract
- •Introduction
- •Materials and Methods
- •Results
- •Discussion
- •Conclusions
- •Acknowledgements
- •References
- •9 Amphibians and Squamate Reptiles from Azokh 1
- •Abstract
- •Introduction
- •Materials and Methods
- •Systematic Descriptions
- •Paleobiogeographical Data
- •Conclusions
- •Acknowledgements
- •References
- •10 Taphonomy and Site Formation of Azokh 1
- •Abstract
- •Introduction
- •Taphonomic Agents
- •Materials and Methods
- •Shape, Size and Fracture
- •Surface Modification Related to Breakage
- •Tool-Induced Surface Modifications
- •Tooth Marks
- •Other Surface Modifications
- •Histology
- •Results
- •Skeletal Element Representation
- •Fossil Size, Shape and Density
- •Surface Modifications
- •Discussion
- •Presence of Humans in Azokh 1 Cave
- •Carnivore Damage
- •Post-Depositional Damage
- •Acknowledgements
- •Supplementary Information
- •References
- •11 Bone Diagenesis at Azokh Caves
- •Abstract
- •Introduction
- •Porosity as a Diagenetic Indicator
- •Bone Diagenesis at Azokh Caves
- •Materials Analyzed
- •Methods
- •Diagenetic Parameters
- •% ‘Collagen’
- •Results and Discussion
- •Azokh 1 Units II–III
- •Azokh 1 Unit Vm
- •Azokh 2
- •Prospects for Molecular Preservation
- •Conclusions
- •Acknowledgements
- •References
- •12 Coprolites, Paleogenomics and Bone Content Analysis
- •Abstract
- •Introduction
- •Materials and Methods
- •Coprolite/Scat Morphometry
- •Bone Observations
- •Chemical Analysis of the Coprolites
- •Paleogenetics and Paleogenomics
- •Results
- •Bone and Coprolite Morphometry
- •Paleogenetic Analysis of the Coprolite
- •Discussion
- •Bone and Coprolite Morphometry
- •Chemical Analyses of the Coprolites
- •Conclusions
- •Acknowledgements
- •References
- •13 Palaeoenvironmental Context of Coprolites and Plant Microfossils from Unit II. Azokh 1
- •Abstract
- •Introduction
- •Environment Around the Cave
- •Materials and Methods
- •Pollen, Phytolith and Diatom Extraction
- •Criteria for the Identification of Phytolith Types
- •Results
- •Diatoms
- •Phytoliths
- •Pollen and Other Microfossils
- •Discussion
- •Conclusions
- •Acknowledgments
- •References
- •14 Charcoal Remains from Azokh 1 Cave: Preliminary Results
- •Abstract
- •Introduction
- •Materials and Methods
- •Results
- •Conclusions
- •Acknowledgments
- •References
- •15 Paleoecology of Azokh 1
- •Abstract
- •Introduction
- •Materials and Methods
- •Habitat Weightings
- •Calculation of Taxonomic Habitat Index (THI)
- •Faunal Bias
- •Results
- •Taphonomy
- •Paleoecology
- •Discussion
- •Evidence for Woodland
- •Evidence for Steppe
- •Conclusions
- •Acknowledgments
- •Species List Tables
- •References
- •16 Appendix: Dating Methods Applied to Azokh Cave Sites
- •Abstract
- •Radiocarbon
- •Uranium Series
- •Amino-acid Racemization
- •Radiocarbon Dating of Samples from the Azokh Cave Complex (Peter Ditchfield)
- •Pretreatment and Measurement
- •Calibration
- •Results and Discussion
- •Introduction
- •Material and Methods
- •Results
- •Conclusions
- •Introduction
- •Laser-ablation Pre-screening
- •Sample Preparation and Measurement
- •Results
- •Conclusions
- •References
- •Index
13 Coprolites and Plant Microfossils |
289 |
Fig. 13.1 The two studied coprolites 5153 (a) and 5246 (b) from Azokh I, Unit II
through modern bioturbation (Murray et al. 2016; Marin-Monfort et al. 2016). During extraction of palaeobotanical remains, a bone fragment was recovered indicating a carnivorous (or omnivorous) diet of the animal that produced the coprolites. Although several other coprolite fragments are available in different layers, only these two were complete and undamaged and were used for plant microfossil extraction.
This paper deals with the extraction of microfossils from the two coprolites in Azokh Cave. With the aim of shedding light on possible environmental conditions that existed during their formation, we investigate the potential of microfossils in the coprolites and discuss them in the context of other fossils and the reconstruction of faunal paleoecology and charcoal that have been found in the deposits from where the coprolites came (Andrews et al. 2016).
Environment Around the Cave
The cave at Azokh (39° 37′ 9.17″ N, 46° 59′ 18.59″ E) is in the Lesser Caucasus at 962 m elevation and the environment is described in this volume (Andrews et al. 2016; Fernán- dez-Jalvo et al. 2016). The rainfall is approximately 600 mm/year, falling mainly in May–June and September– October, while the driest month is January (Republic of Armenia 1999). The faunal contents in the sequence of 300 kyr in the Azokh Cave sedimentary sequence show some variations but are typical of steppe, arid conditions or deciduous woodlands (Andrews et al. 2016). Evidence of the surrounding vegetation in the past can be derived from charcoal in Unit II and Unit Vu, consisting mainly of Prunus (80%) that was probably the most abundant tree species and could have been gathered by humans as firewood while fruits were probably dispersed around the cave (Allué 2016; Andrews et al. 2016).
According to descriptions of present vegetation and plant communities in the Caucasus region it can broadly be divided into three zones: foothill grassland, lower-mountain mixed hardwood forest, and mountain subalpine grassland
(Sharrow 2007). According to global grass distribution maps, this part of the Caucasus consists mainly of species of the Pooideae (c. 300 species), which dominate over other groups like Chloridoideae (17), Paniceae (13), Andropogoneae (6) and Arundinoideae (6). The subfamily Pooideae is the premier group of grasses occupying cool temperate and boreal regions (Cross 1980; Clayton and Renvoize 1986).
Azokh Cave falls in the lower mountain mixed hardwood forest which is generally found at 600–1,100 m elevation (Gulisashvili et al. 1975; Sharrow 2007). At present, most land suitable for farming has been ploughed, and areas suitable for grazing have been grazed. Moderate slopes have often been cleared for use as crop or hay fields, forming large openings in the forest, but areas of forest still exist on steep slopes (Sharrow 2007). The vegetation on the slopes in the vicinity of the cave are currently grassy woodland vegetated by Carpinus, Quercus (probably Q. iberica), Tilia and Fraxinus with an understory of Prunus, Cornus, Corylus, Crataegus and Paliurus spina-christi (Andrews et al. 2016). In the general surroundings Paliurus and Ziziphus is common in “shibliak” (i.e. secondary woodland that develop after forest clearing) (Gabrielian and Fragman-Sapir 2008).
The grasslands of lower elevations once occupied the generally eastern facing foothills and lower slopes of the mountains at about 300–600 m elevation with an annual precipitation of approximately 250–400 mm (Sharrow 2007). Further, cool-season grasses occur with several types of woody species and herbaceous sagebrush in the more xeric areas while shrubs such as buckthorn, hawthorn, and black-wood are found in the more mesic areas.
Materials and Methods
Pollen, Phytolith and Diatom Extraction
The two coprolites (No.’s 5153 = AZ1’08 II-I50#12 and 5246 = AZ1’08 I-H49#4) (Fig. 13.1), which measured 50 × 49 × 33 and 48 × 47 × 30 mm respectively, were sawed in half. One half of each was saved and the other processed for plant microfossil extraction. The studied halves were cleaned by removing the outside 1 to 2 mm layers, which were also saved together with the dust obtained from sawing. They were cleaned further by water to remove dust and then treated in 10% HCl, and cleaned by centrifuging several times using water. Mineral separation was then performed by floating the silica and organic fraction on sodium polytungstate solution (S.G. 2.3) and washing in a centrifuge. Microscope slides were mounted in glycerine jelly and investigated under light microscope, using up to 100× oil