TRENDS IN SUBSISTENCE FROM THE MIDDLE PALEOLITHIC THROUGH

TRENDS IN SUBSISTENCE FROM THE MIDDLE PALEOLITHIC THROUGH
TRENDS IN SUBSISTENCE FROM THE MIDDLE PALEOLITHIC THROUGH
MESOLITHIC AT KLISSOURA CAVE 1 (PELOPONNESE, GREECE)
by
Britt Marie Starkovich
________________________
A Dissertation Submitted to the Faculty of the
SCHOOL OF ANTHROPOLOGY
In Partial Fulfillment of the Requirements
For the Degree of
DOCTOR OF PHILOSOPHY
In the Graduate College
THE UNIVERSITY OF ARIZONA
2011
2
THE UNIVERSITY OF ARIZONA
GRADUATE COLLEGE
As members of the Dissertation Committee, we certify that we have read the dissertation
prepared by Britt Marie Starkovich
entitled Trends in Subsistence from the Middle Paleolithic through Mesolithic at Klissoura
Cave 1 (Peloponnese, Greece)
and recommend that it be accepted as fulfilling the dissertation requirement for the
Degree of Doctor of Philosophy
_______________________________________________________________________
Date:
Dr. Mary C. Stiner
April 26th, 2011
_______________________________________________________________________
Date:
Dr. Steven L. Kuhn
April 26th, 2011
_______________________________________________________________________
Date:
Dr. Vance T. Holliday
April 26th, 2011
_______________________________________________________________________
Date:
Dr. Barnet Pavao-Zuckerman
April 26th, 2011
Final approval and acceptance of this dissertation is contingent upon the candidate’s
submission of the final copies of the dissertation to the Graduate College.
I hereby certify that I have read this dissertation prepared under my direction and recommend
that it be accepted as fulfilling the dissertation requirement.
________________________________________________ Date:
Dissertation Director: Dr. Mary C. Stiner
April 26th, 2011
3
STATEMENT BY AUTHOR
This dissertation has been submitted in partial fulfillment of requirements for an advanced
degree at the University of Arizona and is deposited in the University Library to be made
available to borrowers under rules of the Library.
Brief quotations from this dissertation are allowable without special permission, provided
that accurate acknowledgment of source is made. Requests for permission for extended
quotation from or reproduction of this manuscript in whole or in part may be granted by the
head of the major department or the Dean of the Graduate College when in his or her
judgment the proposed use of the material is in the interests of scholarship. In all other
instances, however, permission must be obtained from the author.
SIGNED: Britt Marie Starkovich
4
ACKNOWLEDGEMENTS
It is difficult to express my thanks and appreciation for all of the people in my life
who have helped make the completion of this project possible. My friends and family have
provided all manner of love and support, and this work is a culmination of all of their efforts.
First of all, thank you to my committee. I managed to find a group of very smart people, who
are also nice to be around. Mary Stiner is a fantastic advisor and has opened doors to me I
never thought imaginable. Steve Kuhn knows everything and more about statistics and stone
tools, and is still on top of pop culture references. Barnet Pavao-Zuckerman patiently
answered bone ID questions in the beginning. Vance Holliday is one of the most down-toearth academics I have ever met, and once scoured west Texas for a BBQ joint with
vegetarian options.
Thanks to the Klissoura Cave 1 team for their excellent work on the site. Margarita
Koumouzelis welcomed me with open arms, and together with Janusz Kozlowski granted me
access to the materials. Takis Karkanas is brilliant, and answers emails with lightening speed.
Thanks also to Piotr Wojtal, Teresa Tomek, Zbigniew Bocheński, Malgorzata Kot, Niko
Thompson, Valery Sitlivy, Krzysztof Sobczyk, Georgia Tsartsidou, Maria Ntinou, and our
Greek workman. Thanks to my colleagues in Greece not affiliated with the Klissoura project.
Sherry Fox is reason enough to work at the Wiener Lab; the faunal collection happens to be a
nice perk. Thanks to Mary Voyatzis and David Romano, and everyone on the Mt. Lykaion
project. Thanks to my undergrad advisors, who helped get me into forager archaeology.
Funding for this study came from a variety of sources, including the NSF IGERT
program, the Reiker Grant and William Shirley Fulton Scholarship funds, the Wiener
Laboratory at the American School for Classical Studies at Athens, an NSF dissertation
improvement grant, and INSTAP, who funded the excavations.
Thanks to my friends, for giving me a mental outlet. This includes, but is not limited
to: Susan Mentzer, Tiina Manne, Nicole Flowers, Mary Good, Natalie Munro, Matt E. Hill,
Rebecca Dean, Amy Clark, Amy Margaris, Shane Miller, Bill Reitze, Katrina Erickson,
Jerome Guynn, Scott St. George, Jake Sutton, Aly Thibodeau, Kevin Anchukaitis, Toby
Ault, Ben Curry, Todd Pitezel, Jess Conroy, Jess Driscoll, Jesse Ballenger, Jenny
Leijonhufvud, Liye Xie, Derek Anderson, Josh Reuther, Lisa Janz, Jon Scholnick, Pat Wrinn,
Matt E. Hill, Joe Beaver, Deanna Grimstead, Jessica Cerezo, Tom Fenn, Dana Drake
Rosenstein, Kelly Jenks, Lauren Jelinek, A.J. Vonarx, Melanie Dedecker Medieros, Katie
MacFarland, Ashley Blythe, Anita Carrasco, Brandon Gabler, Daniela Klöker, Rob Jones,
Lizzie May, Jess Munson, and Ashley Stinnett. Thanks to John Olsen and Barbara Mills. The
tireless efforts of the Anthropology staff have also been critical. Thanks to Ann Samuelson,
Catherine Lehman, Ellen Stamp, Ben Beshaw, Norma Maynard, and especially Cathy Snider.
You are the all the glue that holds us together. Thanks also to my family. Mom, Dad, Ryan,
Christa, aunts, uncles, cousins, Grandma(s) and Grandpa, I love you all so much.
Special thanks, of course, to Mathew Devitt, who has made this project much less
stressful in all ways. Without complaint he has taken care of my beloved pets when I am
doing research, and looked up esoteric sources when I call him from the middle of nowhere.
While I was writing, he also made sure to water and turn me to the sunlight periodically. And
of course, thanks to my furry children (Maggie, Lewis, Tyler, et al.), who keep things in
perspective.
5
TABLE OF CONTENTS
LIST OF FIGURES ...................................................................................................................9
LIST OF TABLES ...................................................................................................................14
ABSTRACT .............................................................................................................................16
CHAPTER 1: INTRODUCTION – QUESTIONS IN PALEOLITHIC ARCHAEOLOGY
AND THEORETICAL BACKGROUND .........................................................................18
INTRODUCTION .......................................................................................................18
THE MIDDLE AND UPPER PALEOLITHIC: BIG QUESTIONS ...........................20
APPLICATIONS FROM EVOLUTIONARY ECOLOGY ........................................29
Prey Choice Model ................................................................................................31
Patch Choice Model ...............................................................................................36
Central Place Foraging Models ..............................................................................39
Predictions of Hominin-Prey Relationships...........................................................48
THE SAMPLE .............................................................................................................49
DISSERTATION STRUCTURE.................................................................................50
CHAPTER 2: KLISSOURA CAVE 1 AND THE GREEK PALEOLITHIC .........................52
INTRODUCTION .......................................................................................................52
NORTHERN GREECE ...............................................................................................54
Asprochaliko ..........................................................................................................54
Theopetra Cave ......................................................................................................58
Kastritsa .................................................................................................................61
Klithi ......................................................................................................................63
Boïla .......................................................................................................................65
Grava ......................................................................................................................66
Sarakenos Cave ......................................................................................................67
Sidari ......................................................................................................................68
Cave of Cyclope .....................................................................................................69
SOUTHERN GREECE ................................................................................................72
Kalamakia Cave .....................................................................................................72
Lakonis I ................................................................................................................74
Apidima..................................................................................................................77
Kephalari Cave.......................................................................................................78
Franchthi Cave .......................................................................................................79
THE LATE MIDDLE PALEOLITHIC THROUGH MESOLITHIC
ARCHAEOLOGICAL SEQUENCE IN GREECE ...............................................85
KLISSOURA GORGE AND KLISSOURA CAVE 1 ................................................88
Stratigraphy and Dating of Klissoura Cave 1 ........................................................92
Stratigraphy ......................................................................................................96
Lithic Assemblages ..............................................................................................100
Botanical Remains ...............................................................................................105
6
TABLE OF CONTENTS - CONTINUED
Features ................................................................................................................106
Ornaments ............................................................................................................108
SUMMARY AND GOALS .......................................................................................110
CHAPTER 3: LATE PLEISTOCENE CLIMATE AND ENVIRONMENT IN THE
EASTERN MEDITERRANEAN ....................................................................................112
INTRODUCTION .....................................................................................................112
ENVIRONMENTAL HISTORY OF THE MEDITERRANEAN: MIS 5A
THROUGH 1 .............................................................................................................113
ENVIRONMENTAL HISTORY OF GREECE: MIS 5A THROUGH 1 .................121
Northern and Central Greece ...............................................................................121
Southern Greece ...................................................................................................127
Environmental Conditions during the Occupation of Klissoura Cave 1 ..............132
BIOTIC COMMUNITIES AND RESOURCE AVAILABILITY ............................135
CONCLUSIONS........................................................................................................139
CHAPTER 4: TAPHONOMIC PROCESSES - BONE ACCUMULATION AND
ATTRITION ....................................................................................................................143
INTRODUCTION .....................................................................................................143
METHODS ................................................................................................................145
RESULTS ..................................................................................................................162
Input Analysis and Fragmentation .......................................................................162
Carnivore Damage and Weathering .....................................................................164
Density-Meditated Attrition .................................................................................168
DISCUSSION AND CONCLUSIONS .....................................................................173
CHAPTER 5: RELATIVE SPECIES ABUNDANCE AND DIVERSITY OF PREY
TYPES .............................................................................................................................177
INTRODUCTION .....................................................................................................177
METHODS ................................................................................................................177
PREY REPRESENTATION IN THE MIDDLE PALEOLITHIC THROUGH
MESOLITHIC LAYERS AT KLISSOURA CAVE 1 ........................................183
PREY DIVERSITY: RICHNESS AND EVENNESS ...............................................193
BIOMASS COMPARISON.......................................................................................200
CONCLUSIONS........................................................................................................204
CHAPTER 6: PREY SELECTION, TRANSPORT DECISIONS AND BUTCHERY
DAMAGE ........................................................................................................................214
INTRODUCTION .....................................................................................................214
BURNING DAMAGE ...............................................................................................216
Burning Damage on Small Game ........................................................................222
Burning Damage on Ungulates ............................................................................224
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TABLE OF CONTENTS – CONTINUED
BUTCHERY DAMAGE AND OSSEOUS TOOLS: CUTS, CONES, IMPACTS,
WORKING, AND OTHER FRACTURES .........................................................229
Butchery Damage on Small Mammals and Carnivores .......................................230
Butchery Damage on Ungulates ..........................................................................233
Worked Bone and Antler .....................................................................................237
BONE PROCESSING ...............................................................................................239
Ungulate Marrow Processing ...............................................................................241
PREY TRANSPORT .................................................................................................245
Utility Indices.......................................................................................................245
Body Part Profiles ................................................................................................252
Small Game Body Part Profiles ...........................................................................253
Ungulate Body Part Profiles ................................................................................259
MORTALITY PROFILES.........................................................................................265
CONCLUSIONS........................................................................................................276
CHAPTER 7: SPATIAL VARIATION WITHIN CULTURAL LAYERS ..........................280
INTRODUCTION .....................................................................................................280
METHODS ................................................................................................................281
RESULTS ..................................................................................................................284
Mesolithic layers 3-5a ..........................................................................................285
Epigravettian layers IIa-d .....................................................................................286
Mediterranean backed bladelet industry layer III’ ...............................................287
Upper Paleolithic (non-Aurignacian) industry layer III” .....................................289
Upper Aurignacian layers IIIb-d ..........................................................................290
Middle Aurignacian layers IIIe-g.........................................................................291
Lower Aurignacian layer IV ................................................................................293
Early Upper Paleolithic or Uluzzian layer V .......................................................296
Middle Paleolithic layer VIII ...............................................................................298
Middle Paleolithic layer X ...................................................................................299
Middle Paleolithic layers XI-XIV ........................................................................299
Middle Paleolithic layers XV-XVII .....................................................................301
Middle Paleolithic layers XVIII-XIX ..................................................................303
Middle Paleolithic layer XXa-b ...........................................................................304
DISCUSSION AND CONCLUSIONS .....................................................................305
CHAPTER 8: KLISSOURA CAVE 1 IN CONTEXT: LATE PLEISTOCENE
SUBSISTENCE CHANGE IN GREECE AND THE MEDITERRANEAN BASIN .....308
INTRODUCTION .....................................................................................................308
KLISSOURA CAVE 1 ..............................................................................................308
Changes in Site Use and Occupation Intensity ....................................................313
RESULTS FROM THE PERSPECTIVE OF EVOLUTIONARY ECOLOGY .......317
Prey Choice ..........................................................................................................317
Central Place Foraging .........................................................................................318
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TABLE OF CONTENTS – CONTINUED
Results on Patch Choice based on Carcass Processing Intensity .........................321
KLISSOURA CAVE 1 WITHIN GREECE AND THE MEDITERRANEAN
BASIN..................................................................................................................324
Intensification and Specialization ........................................................................335
CONCLUSION ..........................................................................................................338
APPENDICES .......................................................................................................................341
Appendix A: Hare survivorship and bone density values (values from Pavao and Stahl
1999) ..........................................................................................................................341
Appendix B: Fallow deer survivorship and bone density values (values from Lyman
1994) ..........................................................................................................................342
Appendix C: Tooth and bone-based MNE values by layer .............................................346
Appendix D: Shaft and end-based MNE values by layer ................................................347
Appendix E: Middle Paleolithic hare elements. Horizontal line indicates lowest structural
density for cervids ......................................................................................................348
Appendix F: NISP and MNE values by layer for all taxon .............................................349
Appendix G: Biomass values for common taxa ..............................................................351
Appendix H: Utility indices for Rangifer, combined by anatomical region (values from
Binford 1978) .............................................................................................................354
Appendix I: Percent MAU for fallow deer by layer ........................................................355
Appendix J: Mean FUI for fallow deer by layer ..............................................................358
Appendix K: Fallow deer anatomical representation.......................................................360
Appendix L: Hare anatomical representation ..................................................................362
Appendix M: Partridge anatomical representation ..........................................................363
Appendix N: Anatomical representation for red deer and ibex in layer IV .....................364
Appendix O: Tooth wear stages for ungulates by layer ...................................................365
Appendix P: Ungulate fusion stages (values from Reitz and Wing 2008 and
Silver 1969) ................................................................................................................367
WORKS CITED ....................................................................................................................370
9
LIST OF FIGURES
Figure 2.1. Map of Greece with Paleolithic and Mesolithic sites mentioned in the text .........55
Figure 2.2. Calibrated radiocarbon dates and ranges of dates for sites mentioned in the text.
Cultural periods are listed above their respective dates. Where radiocarbon dates are
unavailable, placement of cultural periods are estimated based on dated industries in
Greece ................................................................................................................................57
Figure 2.3. Photograph of Klissoura Gorge, facing southwest ................................................89
Figure 2.4. Comparison of modern landmass (grey fill) to LGM shorelines (black line and
white fill, indicating land inundated today) and paleolakes (black fill). Adapted from
Petit-Maire et al (2005) ......................................................................................................90
Figure 2.5. Photograph of Klissoura Cave 1, facing northwest ...............................................91
Figure 2.6. Plan map of Klissoura Cave 1. Adapted from Karkanas (2010) ...........................92
Figure 3.1. Marine oxygen isotope chronology, adapted from Martinson et al. (1987). Bar at
the top indicates period that were warm (w), cold (c) and mild (m), compared to
surrounding periods .........................................................................................................115
Figure 3.2. Simplified schematic of Greek paleoclimate studies mentioned in the text. Dark
bars = warm, wet (forest expansion), grey bars = mild (mixed forest-steppe), white bars =
dry, open steppe. Core locations from studies mentioned in this figure are mapped in
figure 3.3. Bottom axis corresponds to MIS stages in Figure 3.1 ....................................120
Figure 3.3. Map of Greece indicating locations of pollen cores discussed in the text, and in
Figure 3.2 .........................................................................................................................122
Figure 4.1. Metallic blue patina on Middle Paleolithic long bone fragment. From layer
XIX ..................................................................................................................................146
Figure 4.2. (a) Light carbonate fraction on great bustard tibiotarsus. From disturbed zone
(layer 6a). (b) Heavy carbonate fraction on great bustard lumbar vertebrae. From
disturbed zone (layer 6) ...................................................................................................147
Figure 4.3. Chemical weathering on Middle Paleolithic long bone fragment. From layer
XIV ..................................................................................................................................167
Figure 4.4. Proportion of tooth to bone-based MNE for all layers at Klissoura Cave 1. Data
from Table 4.6..................................................................................................................169
Figure 4.5. Proportion of shaft-based and end-based MNE for all layers at Klissoura Cave 1.
Data from Table 4.8 .........................................................................................................172
Figure 5.1. NISP counts for the major taxa at Klissoura Cave 1 by layer. Taxa are in
descending order of average mass ...................................................................................186
Figure 5.2. NISP counts for the major taxa at Klissoura Cave 1, grouped by body class .....187
Figure 5.3. Proportion of small game to ungulate NISP, plotted against time. The relationship
is significant and positive. Data from Table 5.2 ..............................................................189
Figure 5.4. Relative NISP counts for small game animals by layer. White fill indicates small,
slow-moving species, grey or black fill indicates small, fast-moving game ...................190
Figure 5.5. Proportion of small slow to small fast game NISP, plotted against time. The
relationship is significant and negative. Data from Table 5.2 .........................................191
Figure 5.6. NISP counts for carnivore species at Klissoura Cave 1 by cultural layer ...........192
Figure 5.7. Ratio of logNISP to logNtaxa in the Klissoura Cave 1 assemblage. The
relationship is significant and positive. Data from Table 5.3 ..........................................194
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LIST OF FIGURES – CONTINUED
Figure 5.8. Inverse Simpson’s index for all taxa plotted against time. No statistically
significant relationship exists, but there is a general increase in variability between
assemblages in the Upper Paleolithic. Data from Table 5.4 ............................................197
Figure 5.9. Inverse Simpson’s index for all ungulate taxa plotted against time. No statistical
relationship is apparent, though certain Upper Paleolithic and later assemblages are
significantly more even than the other layers. Data from Table 5.4 ................................198
Figure 5.10. Inverse Simpson’s index for small game species plotted against time. No
temporal trend exists, but note the cluster of points that represent higher small game
diversity. The earlier point indicates a more even representation of hares and tortoises,
while the later points indicate layers with an even representation of hares and birds (great
bustard and rock partridge). Data from Table 5.4............................................................199
Figure 5.11. Estimated biomass for each important taxa at Klissoura Cave 1 by layer. Note
that small game biomass is calculated based on MNI, while large game biomass is
calculated using MNE ......................................................................................................201
Figure 5.12. Proportion of small game to ungulate biomass, plotted against time. The
relationship is significant and positive. Data from Table 5.5 ..........................................202
Figure 5.13. Proportion of small slow to small fast game biomass, plotted against time. The
relationship is significant and negative. Data from Table 5.5 .........................................204
Figure 6.1. Frequencies of burned cranial and postcranial specimens at Klissoura Cave 1. In
the Upper Paleolithic layers crania are more commonly burned .....................................228
Figure 6.2. Tortoise carapace fragment with crushing and an impact fracture. From Middle
Paleolithic layer XVI .......................................................................................................232
Figure 6.3. Example of cut marks on medium ungulate remains. Specimen from Middle
Paleolithic layer XII .........................................................................................................236
Figure 6.4. Cut frequencies on fallow deer elements in the Middle Paleolithic layers at
Klissoura Cave 1. Includes elements with n > 10 cuts ....................................................237
Figure 6.5. Worked bone from Middle Paleolithic layer XIV ...............................................238
Figure 6.6. Percent of unopened first, second and third phalanges through the layers at
Klissoura Cave 1. First phalanges, which contain the highest marrow content, are
consistently processed through the layers. Second phalanges were opened more often in
the late Middle Paleolithic and Aurignacian layers. Terminal phalanges were only
intensively processed during the late Middle Paleolithic and Aurignacian layer IV (data
from Table 6.11) ..............................................................................................................244
Figure 6.7. Scatter plots of %MGUI and %MAU values for different layers at Klissoura Cave
1 (following Binford 1978) ..............................................................................................250
Figure 6.8. Mean FUI for each layer, following Broughton (1999). Horizontal line indicates
the average FUI for a complete cervid. Data from Appendix J .......................................252
Figure 6.9. Anatomical profiles for partridges in the Mediterranean backed-bladelet industry
(III’) and upper Aurignacian (IIIb-d) (data from Appendix M) ......................................254
Figure 6.10. Anatomical profiles for hares in the Upper Paleolithic and later layers and
Middle Paleolithic (XI-XIV). Note the lack of foot elements in the UP and Mesolithic
and neck and axial elements in all layers (data from Appendix L) .................................255
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LIST OF FIGURES – CONTINUED
Figure 6.11. Plot of the foot region of hares from Klissoura cave 1 by layer. MAU (or
standardized MNE) values for elements included in the foot are presented in order of
descending structural density from the most dense scan site of each element. The
expected MAU is presented based on the highest MNE value for all skeletal elements for
each layer. Bone density values from Pavao and Stahl (1999) ........................................258
Figure 6.12. Plot of the axial region of hares from Klissoura cave 1 by layer. MAU (or
standardized MNE) values for axial elements are presented in order of descending
structural density from the most dense scan site of each element. The expected MAU is
presented based on the highest MNE value for all skeletal elements for each layer. The
head region is also plotted to indicate that in some layers it is present though other axial
elements are not. Bone density values from Pavao and Stahl (1999) ..............................258
Figure 6.13. Body part profiles for fallow deer in the layers with large sample sizes, ranked
by increasing GUI values (data from Appendix K) .........................................................261
Figure 6.14. Body part profiles for red deer and ibex in lower Aurignacian layer IV, ranked
by increasing GUI values (data from Appendix N) .........................................................262
Figure 6.15. Body part profiles for fallow deer in the layers with large sample sizes,
collapsed into regions with low, medium, high and highest GUI values (from Table
6.16). ................................................................................................................................264
Figure 6.16. Tripolar graph indicating living structure and mortally models in an ungulate
death assemblage .............................................................................................................267
Figure 6.17. Proportions of ungulate age groups exploited at Klissoura Cave 1 during the
Middle and Upper Paleolithic based on tooth eruption and wear. During the Middle
Paleolithic, prime-aged adult animals were targeted. More juveniles were incorporated in
Upper Paleolithic subsistence pursuits. Circles indicate 95% confidence intervals
(following Weaver et al. 2011) (data from Table 6.17) ...................................................270
Figure 6.18. Proportions of fallow deer age groups exploited at Klissoura Cave 1 during the
Middle and Upper Paleolithic based on tooth eruption and wear. During the Middle
Paleolithic, prime-aged adult animals were targeted. Upper Paleolithic patterns are closer
to the living structure of fallow deer. Circles indicate 95% confidence intervals
(following Weaver et al. 2011) (data from Table 6.17) ...................................................270
Figure 6.19. Percentages of juveniles based on five selected elements, based on epiphyseal
fusion, following Stiner (2005), figure 11.12. Selected elements fuse between 12 and 24
months (Kersten 1987). No major differences exist between the ages of juvenile animals
exploited in the Middle and Upper Paleolithic (data from Table 6.17) ...........................272
Figure 6.20. Comparison of fallow deer fetal remains from Klissoura Cave 1 and a modern
stillborn goat. (top) Fetal femur from Upper Paleolithic disturbed zone (6-7a), (bottom)
Fetal humeri from Middle Paleolithic layers (VII) (bottom) and (XVa-XVI) (top) ........274
Figure 6.21. Proportion of fetal to adult fallow deer by MNI. Data from Table 6.19 ...........276
Figure 7.1. Plan maps for Mesolithic (3-5a), NISP = 256. (a) NISP distribution, (b)
frequencies of fresh bone breaks, (c) burning. Darker boxes indicate higher frequencies
of listed parameters. H = unit with hearths ......................................................................286
12
LIST OF FIGURES – CONTINUED
Figure 7.2. Plan maps for Epigravettian (IIa-d), NISP = 67. (a) NISP distribution, (b)
frequencies of fresh bone breaks, (c) burning. Darker boxes indicate higher frequencies of
listed parameters. H = unit with hearths ..........................................................................287
Figure 7.3. Plan maps for Mediterranean backed-bladelet industry (III’), NISP = 1,359. (a)
NISP distribution, (b) frequencies of fresh bone breaks, (c) burning. Darker boxes
indicate higher frequencies of listed parameters. H = unit with hearths..........................288
Figure 7.4. Plan maps for Upper Paleolithic (non-Aurignacian) industry (III”), NISP = 1,617.
(a) NISP distribution, (b) frequencies of fresh bone breaks, (c) burning. Darker boxes
indicate higher frequencies of listed parameters. H = unit with hearths..........................289
Figure 7.5. Plan maps for Aurignacian (IIIb-d), NISP = 1,158. (a) NISP distribution, (b)
frequencies of fresh bone breaks, (c) burning. Darker boxes indicate higher frequencies of
listed parameters. H = unit with hearths ..........................................................................291
Figure 7.6. Plan maps for Aurignacian (IIIe-g), NISP = 1,699. (a) NISP distribution, (b)
frequencies of fresh bone breaks, (c) burning. Darker boxes indicate higher frequencies of
listed parameters. H = unit with hearths ..........................................................................292
Figure 7.7. Plan maps for Aurignacian (IV), NISP = 3,110. (a) NISP distribution, (b)
frequencies of fresh bone breaks, (c) burning. Darker boxes indicate higher frequencies of
listed parameters. H = unit with hearths ..........................................................................293
Figure 7.8. Drafted plan maps in Aurignacian (IV) from 145 to 180 cm below datum by 5cm
cuts. Hearth features are indicated in black and dark grey, limestone rocks in white. The
light grey background represents sedimentary matrix. The shelter feature is apparent in
cuts 145 to 170, where rocks are common and hearths are absent. From Stiner (2010) .295
Figure 7.9. Plan maps for Aurignacian layers (IV) containing the structure, NISP = 2,049. (a)
NISP distribution, (b) frequencies of fresh bone breaks, (c) burning. Darker boxes
indicate higher frequencies of listed parameters. Units containing the structure are
outlined in black...............................................................................................................296
Figure 7.10. Plan maps for Early Upper Paleolithic (Uluzzian V), NISP = 222. (a) NISP
distribution, (b) frequencies of fresh bone breaks, (c) burning. Darker boxes indicate
higher frequencies of listed parameters. H = unit with hearths .......................................297
Figure 7.11. Plan maps for Middle Paleolithic (VIII), NISP = 1,569. (a) NISP distribution,
(b) frequencies of fresh bone breaks, (c) burning. Darker boxes indicate higher
frequencies of listed parameters. H = unit with hearths...................................................298
Figure 7.12. Plan maps for Middle Paleolithic (X), NISP = 241. (a) NISP distribution, (b)
frequencies of fresh bone breaks, (c) burning. Darker boxes indicate higher frequencies of
listed parameters ..............................................................................................................300
Figure 7.13. Plan maps for Middle Paleolithic (XI-XIV), NISP = 1,779. (a) NISP
distribution, (b) frequencies of fresh bone breaks, (c) burning. Darker boxes indicate
higher frequencies of listed parameters ...........................................................................301
Figure 7.14. Plan maps for Middle Paleolithic (XV-XVII), NISP = 2,031. (a) NISP
distribution, (b) frequencies of fresh bone breaks, (c) burning. Darker boxes indicate
higher frequencies of listed parameters ...........................................................................302
13
LIST OF FIGURES – CONTINUED
Figure 7.15. Plan maps for Middle Paleolithic (XVIII-XIX), NISP = 2,199. (a) NISP
distribution, (b) frequencies of fresh bone breaks, (c) burning. Darker boxes indicate
higher frequencies of listed parameters ...........................................................................303
Figure 7.16. Plan maps for Middle Paleolithic (XXa-XXb), NISP = 1,106. (a) NISP
distribution, (b) frequencies of fresh bone breaks, (c) burning, (d) medium ungulate
specimens by body region. Darker boxes indicate higher frequencies of listed
Parameters ........................................................................................................................305
Figure 8.1. Illustration of changes through time in ungulate diversity, occupation intensity
and small game use at Klissoura Cave 1 ..........................................................................315
Figure 8.2. Map of the four quadrants of the Mediterranean. Shaded areas indicate
Mediterranean ecosystem. From Blondel and Aronson (1999) .......................................324
14
LIST OF TABLES
Table 2.1. Stratigraphic layers for Klissoura Cave 1 and corresponding cultural units based
on stone tool types. Layer groupings are used throughout the dissertation for analyzing
faunal remains. From Kaczanowksa et al. 2010, Karkanas 2010, personal communication
2009, Kuhn et al. 2010 .......................................................................................................93
Table 2.2. Radiocarbon dates from Klissoura Cave 1. From Kuhn et al. (2010) ....................95
Table 3.1. Taxa expected in the vicinity of Klissoura Cave 1 by MIS stage. Note that the MIS
stage of the lower layers are unknown, which is indicated by "/" and both options are
included. MIS 3 is highly variable and local reconstructions are available for the UP
layers but not the MP. P = present, A = absent, ? = unsure .............................................133
Table 4.1. Fallow deer metatarsal NISP counts and NISP:MNE ratios that include anterior
groove fragments and corrected counts that exclude anterior groove fragments, compared
to metacarpal NISP counts and ratios. Metacarpals were used for comparison because
their structural density is similar and they are equally recognizable in an assemblage as
are metatarsals. NISP counts for metatarsals exclude anterior grooves for taxonomic
frequencies because they over-represent fallow deer frequencies. Only Middle Paleolithic
layers are depicted because they are the most extreme, but cervid metatarsal anterior
grooves (and other overly-recognizable fragments such as unidentifiable tooth and
tortoise shell fragments) are excluded in all layers ..........................................................150
Table 4.2. Rates of faunal input (or faunal density) by layer ................................................151
Table 4.3. Mean fragment length for different body classes, by cultural layer .....................164
Table 4.4. MNE:NISP index values for medium ungulate cranial elements and all elements in
the Klissoura Cave 1 assemblage. *Note small sample sizes ..........................................165
Table 4.5. Frequencies of gnawing and weathering damage on faunal specimens by cultural
layer..................................................................................................................................166
Table 4.6. Tooth and bone-based MNE for all ungulates by layer ........................................169
Table 4.7. Spearman’s rank-order correlation between percent survivorship and bone density
values for European hare and fallow deer/medium ungulate remains. Data from
Appendices A and B. *Significant values........................................................................171
Table 4.8. End and shaft-based MNE values for medium ungulate long bones by cultural
layer. Elements include humerus, radius, ulna, metacarpal, femur, tibia and
metatarsal .........................................................................................................................172
Table 5.1. Taxa included in each body class category and their respective weight ranges and
predicted ranking. Mass values from Nowack (1999) and Silva and Downing (1995) ...179
Table 5.2. Proportion of NISP counts per layer for small game: large game (ungulates), and
small slow: small fast-moving prey types. *Small samples excluded from Figure 5.3 ...188
Table 5.3. LogNISP and logN-taxa values for all species, ungulates, and small game
Species .............................................................................................................................195
Table 5.4. Inverse Simpson’s index per layer for all taxa, ungulates, and small game species.
*Small sample sizes excluded from Figure 5.10 .............................................................196
Table 5.5. Proportion of biomass values per layer for small game: large game (ungulates),
and small slow: small fast-moving prey types. *Small samples excluded from Figure
5.13...................................................................................................................................203
15
LIST OF TABLES – CONTINUED
Table 5.6. Taxa found in Klissoura Cave 1 by cultural layer, with predictions from Table 3.1
in parentheses. P = present, A = absent, U = uncommon (NISP < 5). Grey shading
indicates deviations from predictions. Roe deer and chamois are excluded because of
small samples overall. *Epigravettian NISP = 67 ...........................................................206
Table 6.1. Burn frequency and degree for different classes of animals at Klissoura Cave 1 by
layer..................................................................................................................................218
Table 6.2. Burn frequencies for tortoise shells in layers with larger sample sizes ................223
Table 6.3. Mean length for burned and unburned identifiable specimens by layer ...............224
Table 6.4. Burn frequency and degree for cranial and postcranial elements by layer for
medium ungulates. *Small samples not included in Figure 6.1 ......................................226
Table 6.5. Butchery damage on small game at Klissoura Cave 1 by layer ............................231
Table 6.6. Transverse fractures on small animal limb bones (humerus, radius, femur and
tibia/tibiotarsus) by layer .................................................................................................233
Table 6.7. Butchery damage on medium and large ungulate remains. *Note small sample
sizes ..................................................................................................................................234
Table 6.8. Cut marks on medium ungulate elements in Middle Paleolithic layers ...............236
Table 6.9. Counts of all antler and worked antler by layer, and corresponding ungulate NISP
counts for comparison ......................................................................................................239
Table 6.10. Percent of medium ungulate limb bones not opened prior to discard, by layer .242
Table 6.11. Percent of unopened medium ungulate phalanges for layers with MNE values
larger than 10 ...................................................................................................................243
Table 6.12. Pearson’s correlation values between Binford’s (1978) utility indices. * Indicates
significant relationships ...................................................................................................247
Table 6.13. Spearman’s rank-order correlation values for fallow deer %MAU and caribou
utility indices (from Binford 1978). *Statistically significant relationships....................249
Table 6.14. K-S tests for hare body part profiles by layer. *Asymp. Sig. > 0.05, data do not
differ from a uniform distribution ....................................................................................255
Table 6.15. K-S tests for ungulate body part profiles by layer. *Asymp. Sig. > 0.05, data do
not differ from a uniform distribution ..............................................................................262
Table 6.16. Cervid anatomical regions (following Stiner 1991), ranked in order of GUI
values derived for caribou (from Binford 1978) ..............................................................263
Table 6.17. Age distribution for ungulates in the Klissoura Cave 1 assemblages, based on
tooth eruption data (see Appendix O for more detail) .....................................................269
Table 6.18. Fusion schedules for five elements for fallow deer. Note radii are excluded from
Figure 6.19 because of small samples in the Upper Paleolithic layers ............................273
Table 6.19. Fetal elements identifiable to species by layer ...................................................275
16
ABSTRACT
This study presents an analysis of the zooarchaeological remains from Klissoura Cave
1, a Middle Paleolithic through Mesolithic site in Peloponnese, Greece. Changes in
subsistence patterns are evaluated across a long sequence (ca. 80,000-10,000 BP) against a
backdrop of environmental change. Results are interpreted using models from evolutionary
ecology, specifically prey choice, central place foraging, and patch choice models. Two
major trends are apparent in the series. One is a decline in the exploitation of high-ranked
ungulate species with an overall increase in lower-ranked small game animals. The second is
an increase in low-ranked small, fast-fast moving animals (e.g., hares and partridges) at the
expense of higher-ranked small, slow-moving animals (e.g., tortoises). These changes cannot
be accounted for by environmental shifts alone, though shifts in ungulate diversity likely
track the expansion and contraction of plant communities. The increase in use of low-ranked
prey indicates human population growth and demographic pressure in southern Greece
during the late Pleistocene and early Holocene. In addition to these overarching trends, there
are changes in site use during the sequence. In the Middle Paleolithic, foragers used
Klissoura Cave 1 more during the winter and overwhelmingly hunted prime-aged adult
animals, maternal herds of fallow deer in particular. In the Upper Paleolithic and later
periods, the site continued to be used during the winter, in addition to other times of year, but
the mortality profiles reflect a natural fallow deer herd structure. There was an intense period
of occupation during the Aurignacian period. This is evidenced by numerous clay-lined
hearth features, a possible rock-lined structure, and increases in ornaments, as well as
abundant lithic and faunal materials. The ungulate faunas are particularly rich during this
17
period, but there is evidence of resource intensification based on increased bone marrow
processing and the transport of marrow-rich elements to the site. After this period there was a
gradual decline in site use through the end of the Upper Paleolithic and into the Mesolithic,
though the exploitation of low-ranked resources (e.g., small, fast-moving game) indicates
that populations were on the rise in there region as a whole.
18
CHAPTER 1: INTRODUCTION - QUESTIONS IN PALEOLITHIC
ARCHAEOLOGY AND THEORETICAL BACKGROUND
INTRODUCTION
On a global scale, the Middle and Upper Paleolithic were extremely dynamic periods,
and change accelerated over time as hominin populations expanded into new regions and
local cultural diversity increased. Human population densities fluctuated dynamically as
anatomically modern humans left Africa and pushed into regions already occupied by
Eurasian hominins. Stone tool technologies became more complex and locally distinct.
Hominin subsistence strategies shifted in response to environmental change, tighter packing
of humans on the landscape, and technological innovations that allowed for the safer hunting
of large game (e.g., projectile weaponry) and the more efficient exploitation of small game
(e.g., snares, nets and fish hooks). Durable artistic and symbolic expression appeared in
Eurasia and became ubiquitous during the Upper Paleolithic, including shell and tooth
ornaments, ochre-stained tools and bone, figurines, and eventually cave paintings. In general
it was a time of tremendous social and cultural upheaval. As archaeologists have tried to
make sense of variation within and between the Middle and Upper Paleolithic, their jobs are
complicated by an archaeological record that is often geographically and temporally patchy,
dating techniques with low levels of precision, and a lack of hominin skeletons that make it
difficult to assign stone tool industries to specific populations.
It is unrealistic to assume that any work, no matter how ambitious, could solve all of
these problems, but every well-designed research project can add to our understanding of the
final chapters of the evolutionary history of human foragers. This introduction serves as a
19
backdrop for a discussion of human dietary change across the late Pleistocene and early
Holocene from Klissoura Cave 1, a small rock shelter in southern Greece. The archaeological
record at Klissoura Cave 1 is important for several reasons. First is the overall lack of long,
intact archaeological sequences in Greece, which is discussed in detail in Chapter 2. Second,
is the importance of Greece in general to our understanding of hominin lifeways and
population movements during the late Pleistocene. The geographic significance of Greece
cannot be overemphasized. Along with Iberia and Italy, it is one of three major European
peninsulas, and there is ample evidence that some of these regions served as refugia during
the Pleistocene for plant, animal, and hominin populations (see below). Further, its location
in eastern Europe makes Greece key in understanding the western expansion of modern
human populations.
The Klissoura Cave 1 record is equally important for understanding variation within
cultural periods as it is for examining broad changes between periods. However, many
changes concerning Paleolithic archaeology are framed in terms of understanding the socalled transition between the Middle and Upper Paleolithic. As such, I review some
arguments and debates pertaining to difference between the two periods, and the difficulties
of characterizing “transitional” archaeological phases in Europe. Often, these discussions
revolve around the behavioral dichotomy (or not) between Neandertals and anatomically
modern humans, whose cultural remains are presumably both represented at Klissoura Cave
1. An absence of hominin skeletal remains at the site, however, makes this dichotomy
difficult to explore. Therefore, this study focuses on changes in diet and subsistence practices
within and between cultural phases, regardless of whether they are associated with changes in
hominin populations or lithic technologies. These changes are explored throughout this
20
dissertation using theoretical tools from evolutionary ecology, which are expanded upon in
the latter part of this chapter.
THE MIDDLE AND UPPER PALEOLITHIC: BIG QUESTIONS
Many complex questions surround attempts to understand the Middle and Upper
Paleolithic in Eurasia, most of which focus on the nature and meaning of the transition
between the two periods. The nature of the transition is contentious and difficult to
understand because there was no “moment” that indicates the shift to behavioral and cultural
modernity (Clark 2002; Hovers and Belfer-Cohen 2006; McBrearty and Brooks 2000). The
general timing of the transition seems to have varied, with evidence of behavioral modernity
appearing earlier in Africa and later in southwest Asia and Europe (see McBrearty and
Brooks 2000 and references therein). Another question seeks to determine exactly who the
Neandertals were, a local population of Homo sapiens that contributed, albeit minimally, to
the modern human gene pool (Ahern, et al. 2004; Clark 2002; Trinkhaus 2007; Zilhão 2006),
or a behaviorally and technologically stagnant “other” that sometimes relied on anatomically
modern humans for cultural innovations (d'Errico, et al. 1998; Mellars 1973, 1989, 1991).
The abilities of Neandertals to hunt efficiently, create art, and innovate stone tool
technologies have been widely questioned (Mellars 1973, 1989, 1991). The extent to which
populations of modern Homo sapiens from Africa interacted with European Neandertals is
also unclear, whether there was interbreeding (but see Green, et al. 2010), reluctant tolerance,
the transmission of cultural knowledge, or outright hostility. It is also not known exactly how
long Neandertal populations persisted in the refugia of the various southern European
peninsulas (Carbonell, et al. 2000; Conard and Bolus 2003; d'Errico and Sanchez Goñi 2003;
21
d'Errico, et al. 1998; Finlayson, et al. 2006; Kuhn and Biette 2000; Tzedakis, et al. 2007;
Walker, et al. 2008; Zilhão and d'Errico 1999; Zilhão and Pettitt 2006). What is apparent,
however, is that human population densities increased around the world within the interval
between the late Middle Paleolithic and the mid-Upper Paleolithic (e.g., Lahr and Foley
2003), and a range of cultural adaptations appeared, at least partly in response to population
growth against a backdrop of climatic change. The current evidence for the emergence of
durable forms of artistic expression, new stone tool chronologies, bioarchaeological data, and
subsistence change are reviewed briefly below.
One of the hallmarks of behavioral modernity is arguably the emergence of symbolic
art. The lack of artistic expression at European Middle Paleolithic sites was long considered
as evidence for cognitive and social differences between anatomically modern humans and
Neandertal populations in particular (Mellars 1973, 1989, 1991). Ochre and black manganese
occur throughout the Middle Paleolithic record, the uses of which are unknown. Evidence of
artistic expression is very rare or absent in Middle Paleolithic contexts, though shell
ornaments, ochre-stained artifacts and lumps of ochre are reported in a handful of sites
outside of Africa, from Spain to the Levant and dating to between 135,000 and 50,000 BP
(Bar-Yosef Mayer, et al. 2009; Vanhaeren, et al. 2006; Zilhão, et al. 2010). All early contexts
containing Middle Paleolithic artistic expression, except those reported from Spain, are
associated with anatomically modern humans. There is a difference between the sporadic
artistic expression claimed for some Middle Paleolithic cases and its ubiquity and
unmistakable form in Upper Paleolithic and later contexts (e.g. Kuhn and Stiner 2007; Kuhn,
et al. 2001; Stiner 2010). It is no longer fair to assume that Neandertals and other Middle
22
Paleolithic populations were cognitively incapable of creating symbolic expression, but the
lack of widespread durable examples is very difficult to explain or ignore.
The chronology of the transition from the Middle to Upper Paleolithic varies
regionally, beginning earlier in southwest Asia and occurring the latest in western Europe.
The transition began between roughly 50,000 and 45,000 BP in the Levant (Bar-Yosef 2000;
Bar-Yosef 2007), and gradually spread through Eurasia. The transition in the Caucasus, for
example, occurs between about 35,000 and 34,000 BP (Adler, et al. 2006; Meshveliani, et al.
2004). The Middle Paleolithic persisted in Eastern Europe until between 47,000 and 40,000
(Kozlowski 2000; Kozlowski 2007), before the Eastern European Aurignacian became
widespread. The transition to the Aurignacian occurred after 36,500 BP in France and Spain
(d'Errico, et al. 1998; Zilhão and d'Errico 1999).
Some areas of Europe, such as the southern Iberian and Italian peninsulas may have
served as refugia for Neandertal populations (Carbonell, et al. 2000; Kuhn and Biette 2000).
Late-persisting Neandertals still manufacturing Mousterian technology existed in southern
Iberia as late as 28,000-30,000 BP (Carbonell, et al. 2000; d'Errico and Sanchez Goñi 2003;
d'Errico, et al. 1998; Finlayson, et al. 2006; Zilhão and d'Errico 1999; Zilhão and Pettitt
2006). In Italy, it was initially thought that there was about a five thousand year difference
between the appearance of the Aurignacian industry in the north and south, with Neandertal
populations persisting until 32,500 BP (Kuhn and Biette 2000). However, recent re-dating of
layers at Grotta di Fumane now suggest that transitional Uluzzian and Proto Aurignacian
industries were deposited before 39,900 BP (Higham, et al. 2009; Peresani, et al. 2008). In
fact, Higham et al. (2009) postulate that many sites dating to between 35,000 and 45,000 BP
23
in Italy are actually much more contemporaneous than traditional AMS dating techniques
indicate.
One of the major issues in European Paleolithic archaeology is the extent to which
anatomically modern humans and Neandertal populations interacted, and how these
interactions, or lack thereof, are reflected in early Upper Paleolithic stone tool industries.
Some scholars argue that in France, Neandertal and anatomically modern human populations
overlapped between about 38,000 and 33/34,000 BP. During this time Neandertals
presumably “borrowed” technological attributes from anatomically modern humans,
culminating in certain early Upper Paleolithic stone tool industries such as the
Châtelperronian (Mellars 1999). Other authors reject this claim (Clark 2002; d'Errico, et al.
1998; Zilhão and d'Errico 1999). Clark (2002) points out that the Aurignacian industry is
primarily based on blade production, while the twenty plus regional Early Upper Paleolithic
industries do not necessarily include blade production. D’Errico and colleagues (1998; Zilhão
and d'Errico 1999) also disagree with Mellars (1999) about Neandertal “enculturation.” They
present a re-evaluation of Grotte du Renne in order to establish the cultural originality of the
Châtelperronian Neandertal occupants of the site. They point to a stratigraphically sound
Châtelperronian layer beneath the Aurignacian occupation, and debitage from bone tool and
ornament manufacture in the former (but see Higham, et al. 2010 for recent dates that
indicate the bone tools and ornaments are actually intrusive). D’Errico and colleagues (1998;
Zilhão and d'Errico 1999) discuss differences in the reduction sequences, blank selection and
tool manufacture between the two industries. Most notably, unlike the Aurignacian, the
Châtelperronian contains no large-scale bladelet production. In general, degree of interaction
is difficult to assess based on archaeological evidence because of the potential for
24
stratigraphic mixing, and unclear implications from interstratifications of cultural layers even
in the case that they are found intact (e.g., did one population replace another, or did they
simply use a site after it was abandoned by a different group).
The bioarchaeological record, including hominin fossils and paleoDNA, have the
potential to determine if there was any gene flow between Neandertal and modern human
populations. Mitochondrial DNA studies have consistently indicated that there was little or
no admixture between modern human and Neandertal populations (Briggs, et al. 2009;
Krings, et al. 1997; Ovchinnikov, et al. 2000; Serre, et al. 2004). More recent studies of long
sequences of nuclear DNA, however, suggest that small amounts of Neandertal DNA (1-4%)
are present in the genomes of modern Eurasian populations (Green, et al. 2010). These genes
are not present in modern African groups, which indicates that gene flow occurred after the
initial divergence between modern human populations in Africa and European Neandertals,
or that the lineages went extinct in Africa.
Human fossils associated with “transitional” or Early Upper Paleolithic industries
may indicate the authors of these stone tool types. A problem with the skeletal data is that
early Upper Paleolithic skeletons are extremely rare and it is difficult to construct a model of
what contemporary anatomically modern human populations looked like, even in Africa. A
small handful of European skeletons from well-dated contexts have traits consistent with
both Neandertal populations and the few contemporary modern humans (Duarte, et al. 1999;
Trinkhaus 2007; Trinkhaus, Milota, et al. 2003; Trinkhaus, Moldovan, et al. 2003; Wild
2005; Zilhão 2006). This largely supports the nuclear DNA data results, namely that there
was a small amount of gene flow between the two populations.
25
The paucity of hominin skeletal remains also makes it difficult to determine the
manufacturers of key Early Upper Paleolithic stone tool industries (e.g., the Châtelperronian,
Uluzzian and Szeletian), since stone tools and human remains are rarely found in direct
association with one another. A reanalysis of the hominin remains from Vindija Cave in
Croatia identifies Neandertal remains with Upper Paleolithic lithic and bone technology,
though there is potential stratigraphic mixing in the associated layers (Ahern, et al. 2004).
Neandertal remains are associated with the late Micoquian in Germany (Schmitz, et al.
2002), the French Châtelperronian (Hublin, et al. 1996, but see Bar-Yosef and Bordes 2010)
and possibly the Italian Uluzzian (Churchill and Smith 2000, but see Riel-Salvatore 2009).
There is consensus among many archaeologists that the Aurignacian industry is connected
with anatomically modern humans, though bioarchaeological remains from this period are
very rare (Gambier 1989; Mellars 1989, 1991; Stringer, et al. 1984; Svoboda, et al. 2002).
Early questions concerning Paleolithic subsistence strategies have focused on whether
Neandertals were successful hunters or obligate scavengers. Though this debate is fairly well
exhausted at this point, it has important implications if Neandertals were similar to Africanorigin modern human populations in terms of foraging ecology, social structure, and
language abilities. Differences in these capacities may explain why modern humans
eventually out-competed Neandertals.
Binford (1984b; 1985; 1988) was a major proponent of middle and early late
Pleistocene hominins as scavengers based on faunal assemblages from Klasies River Mouth
Cave in South Africa and Grotte Vaufrey in France. He interpreted the French assemblage as
originating from carnivore kills that were later scavenged by humans and brought to the site
for processing (Binford 1988). The African remains were dominated by head and foot
26
elements which were thought to be indicative of scavenging, though some small game
hunting was also noted (Binford 1984b). Binford’s assertions drew early criticism, and many
subsequent studies have disagreed with his conclusions (Chase 1989; Grayson and Delpech
1994; Marean and Kim 1998; Speth and Tchernov 1998; Stiner 1994).
In a survey of European Paleolithic sites, Chase (1989) finds few differences in
hunting practices between Middle and Upper Paleolithic hominins; Neandertal assemblages
indicate foresight in their exploitation of migratory ungulate herds and occasional
specialization in one large game species. Grayson and Delpech (1994) present a taphonomic
reanalysis of the fauna from Grotte Vaufrey and find that Binford’s (1988) conclusion that
the Middle Paleolithic inhabitants of the site primarily scavenged large game is not
supported. In a comprehensive taphonomic and economic analysis of faunal remains from
Middle Paleolithic sites in Italy, Stiner (1994) examines Neandertal subsistence strategies as
compared to other top-tier late Pleistocene carnivores. She determines that based on prey
choice, body part representation and age structure analysis, Neandertals were successful
hunters of prime-aged adult large game, as well as occasional, opportunistic scavengers, not
unlike many modern foragers (e.g. O'Connell, et al. 1988). Similar conclusions, that
Neandertals and even earlier Eurasian hominins targeted prime-aged adult animals are noted
in Middle and Lower Paleolithic sites in Israel (Speth and Clark 2006; Speth and Tchernov
1998; Stiner 2005; Stiner, et al. 2009), the Republic of Georgia (Adler, et al. 2006), and
northern Europe (Gaudzinski 1995; Gaudzinski and Roebroeks 2000). There is also evidence
that Neandertal groups in Gibraltar and Italy occasionally exploited marine mammals in
addition to marine mollusks (Stiner 1994; Stringer, et al. 2008). As these studies suggest,
27
Neandertals were clearly successful hunters of large game, and the hunting/scavenging
debate has been laid to rest.
Another hunting behavior traditionally considered as a hallmark of the Upper
Paleolithic is specialized hunting a single large game species. This idea, proposed by Mellars
(1973; 1989; 1991) was based on some instances of monospecific faunas such as reindeer
assemblages in Paleolithic sites in western France. Such hunting strategies were thought to
have a link to cognition as an indicator of the capacity for planning, forethought and logistics
that accompanies predicting the movements of gregarious taxa (Mellars 1989), though
natural diversity and seasonality also play a part. Since specialization was thought to be
unique to the Upper Paleolithic, Neandertals and other Middle Paleolithic hominins were
apparently lacking these high-level cognitive functions.
Monospecific faunas are documented at many Upper Paleolithic sites, particularly
those that concern ibex hunting (e.g. Gamble 1997; Gamble 1999; Phoca-Cosmetatou 2004a;
Straus 1987), but it turns out that they are not unique to the Upper Paleolithic.
"Specialization" as defined by the presence of monotypic assemblages occurs as early as the
Middle Paleolithic (e.g. Costamagno, et al. 2006; Gaudzinski 1995, 2006) and represents a
range of hunting strategies, including mass kills, and long-term periodic exploitation of herd
and solitary animals. In the Middle and Upper Paleolithic of southern France, certain species
were targeted during seasonal migrations (Costamagno 2004; Costamagno, et al. 2006;
Surmely, et al. 2003).
Examining small game exploitation patterns during the Paleolithic, Stiner and
colleagues (Stiner 2001; Stiner and Munro 2002; Stiner, et al. 2000) show that the relative
proportions of small to large game exploited during the Middle and Upper Paleolithic periods
28
in Italy and Israel did not uniformly change, though the kinds of small game exploited
drastically shifted. In the Middle Paleolithic, small, slow-moving game species such as
tortoises and limpets were exploited, while in the Early Upper Paleolithic, game birds and
small mammals became proportionally more common. By the Epipaleolithic, lagomorph
abundance was especially high (Stiner, et al. 2000). Additionally, Stiner et al. (2000) present
data on size diminution in limpets from Italian sites and tortoises from sites in Israel, which
indicates increased levels of pressure on these slow-reproducing species as early as 44,000
BP. A similar situation may have occurred in Turkey at Üçağızlı Caves I and II, where
ungulates dominate the Middle and Upper Paleolithic layers, though they decline by the
Epipaleolithic. Small, slow-moving taxa are important in the Middle Paleolithic, and again in
the Ahmarian (Stiner 2009; Stiner, et al. 2002). By the Epipaleolithic, small fast-moving prey
vastly increases in dietary importance. This pattern is also beginning to emerge in southern
Greece, particularly by the Epipaleolithic and Mesolithic periods (Starkovich 2009;
Starkovich and Stiner 2010; Stiner and Munro 2011). Small game was less important in
southern France and areas to the north, as compared to the eastern Mediterranean
(Costamagno 2004; Costamagno, et al. 2006; Surmely, et al. 2003).
Clearly, zooarchaeology has much to tell us about the nature of hunting patterns and
human adaptations across the late Pleistocene. Based on the studies reviewed above, drastic
differences are not expected between the faunal assemblages generated by Neandertals and
anatomically modern humans when it comes to planning or organizing hunting activities.
Where there are differences, explanations related to environmentally driven changes in prey
structure, human hunting pressures, or changes in site use or occupation intensity are
evaluated, as opposed to the assumption that there was a cognitive difference between
29
Middle and Upper Paleolithic hominins. Faunal studies are more instructive in understanding
other kinds of changes that occurred during the Paleolithic, such as responses to local
environmental shifts or rising human population densities, either of which can stress prey
populations. The Klissoura Cave 1 occupation spans the late Pleistocene and early Holocene,
and includes Middle Paleolithic through Mesolithic cultural layers. Understanding variation
within as well as between these occupations are central to this study. Theoretical models
from evolutionary ecology are important for formulating predictions pertaining to changes in
hunting and processing strategies through the occupation of the site.
APPLICATIONS FROM EVOLUTIONARY ECOLOGY
Human dietary change may reflect a variety of internal and external factors, from
climatic shifts to population growth, changes in landscape use and site occupation intensity,
or cultural preferences. Evolutionary ecology is a useful paradigm for framing questions
about shifts in game use (see review in Bird and O'Connell 2006; Lupo 2007). Foraging
models in particular are based on the idea that organisms maximize their fitness by making
decisions that optimize their foraging efficiency and reproductive success within a given set
of dynamic environmental circumstances (Pianka 2000). The paradigm is associated with a
series of theories and behavioral models (sometimes called optimal foraging models) that
explain foraging decisions: these include the diet breadth or prey choice model (Pianka 2000;
Stephens and Krebs 1986), patch choice models (MacArthur and Pianka 1966), and central
place foraging models (Orians and Pearson 1979; Schoener 1979). Other models are also
employed by evolutionary ecologists (e.g., costly signaling theory and tolerated theft) but are
30
not discussed further as the three models mentioned previously are more readily applied to
the Paleolithic archaeological record.
Many criticisms have been leveled at evolutionary ecology and its application to
human foragers in the ethnographic and archaeological records (e.g. Dwyer 1986; Joseph
2000; Martin 1983; Pyke 1984). The critiques offer some valuable and often nuanced points;
the major misgivings seem to focus on 1) a discomfort with applying mathematical models to
human behavior, because they are too simplistic or mechanistic and 2) practical matters such
as defining the currency used to test optimality models. Some allies of evolutionary ecology
also point out that due to the nature of the archaeological record, foraging models tend to
focus on the faunal portion of the diet and ignore the vegetal component (Broughton and
Grayson 1993; Grayson and Cannon 1999). Hill et al. (1987) also point out some of the
difficulties of testing optimal foraging models. Specifically, an ethnoarchaeologist cannot
coerce foragers to exploit low-ranked resources, so it is impossible to quantify their return
rate. Because of this, it is unclear if certain items are excluded from the diet because of their
low nutritional return, or for other, possibly cultural reasons. These criticisms are addressed
extensively elsewhere (Broughton and O'Connell 1999; Smith 1991, 2000; Smith and
Winterhalder 1985; Winterhalder 2002).
The existence of potential problems with some aspects of optimal foraging models
does not render them useless to anthropologists. The thought of humans conforming to
simple mathematical models is may be unsettling, as such models may seem to deny us free
will or cultural preference. More is learned, however, from deviations from expectations put
forth by evolutionary ecology and optimal foraging models than from cases that fit
predictions. The second criticism addresses the currency and goals used to test optimality
31
models. Typically, evolutionary ecologists focus on kilocalories as the unit of measurement
of cost and gain in foraging pursuits, but other nutritional factors, such as complete protein or
fat yields, may actually be more important. Further, goals may change depending on who is
doing the foraging, and if they are responding to specific circumstances such as provisioning
offspring. In general, these issues are addressed in the ethnographic literature (e.g. Bird and
Bliege Bird 1997; Hill and Hawkes 1983; Hill, et al. 1987; Kelly 1995; Smith 1991), but they
are difficult to test in the archaeological record because there are so many more unknowns.
Kilocalories, and in most cases prey mass (see below) are used as proxy measures of
nutritional value in most studies, and it is acknowledged that they are only estimations. As
far as paying more attention to the meat component of archaeological diets, this is true in the
same way that technological studies focus more on stone than organic tools in most
instances. If organic remains, such as plant foods or perishable technologies preserve at a
site, they should absolutely be included in considerations of diet breadth or technological
adaptations. But the absence of such evidence does not mean that we cannot use models from
evolutionary ecology to help understand exploitation of faunal resources.
Prey Choice Model
The diet breadth, or prey choice model, is particularly helpful for understanding
dietary change among foragers (Pianka 2000; Stephens and Krebs 1986). The prey choice
model assumes that resources are encountered randomly and sequentially on a landscape.
Prey items are ranked according to their return rate, which is a measure of the caloric or
nutritional value of a resource per unit of time invested in obtaining it, and includes both
search and post-encounter handling costs (Pianka 2000; Stephens and Krebs 1986). Foragers
32
attempt to maximize their net caloric return by preferring prey with the highest net returns.
Following the prey choice model, foragers are always expected to pursue high-ranked prey
whenever they are encountered on the landscape. Further, the decision to pursue low-ranked
species is irrespective of their abundance in the environment; rather foragers only add lowranked species to the diet as high-ranked resources become scarce (Emlen 1966; MacArthur
and Pianka 1966; Pianka 2000; Stephens and Krebs 1986).
Other variables can influence prey selection among humans, such as composition of
the foraging party (Bird and Bliege Bird 2000; Hill, et al. 1987; Hurtado, et al. 1985; Jochim
1988; Lupo and Schmitt 2002, 2005), prestige (Hawkes 1990, 1991; Hawkes and Bliege Bird
2002; Smith 2004), or cultural taboos or preferences for certain resources. Since the
archaeological record represents a palimpsest of foraging events, it is assumed for the
purposes of this study that the remains at Klissoura Cave 1 are an average of the prey items
exploited by foraging groups of variable composition (e.g., men and women) and that any
temporal trend reflects changes in the foraging strategies of the entire group.
Many ethnographic studies have employed prey choice models in order to examine
human hunting decisions (Hames and Vickers 1982; Hawkes, et al. 1982; Hawkes and
O'Connell 1981, 1985; Hill and Hawkes 1983; Hill, et al. 1987; Smith 1991; Winterhalder
1981). Hawkes and O’Connell (1981) provide an early application of the prey choice model
to human foragers, in this case the gathering of tree and grass seeds by Alyawara women in
Australia. By taking into account collection and processing costs, they determine that
workloads for many foragers are actually considerably higher than was initially reported by
Lee (1979) in his work with the !Kung San, who were presented as the model case for the
“affluent forager.” Indeed, including processing costs in the calculation of foraging
33
efficiency drastically decreases the return rate of mongongo nuts, a key dietary resource of
the !Kung (Hawkes and O'Connell 1981).
Prey choice models are utilized in a series of papers that focus on the Aché, a group
of semi-settled hunter-gatherers in eastern Paraguay (Hawkes, et al. 1982; Hill and Hawkes
1983; Hill, et al. 1987). In this case animals, particularly large-bodied prey, tend to be ranked
higher than plant resources, especially plant foods that require processing (Hawkes, et al.
1982; Hill, et al. 1987). Further, Hawkes, Hill and colleagues find that the resources during
the study period yielded a return higher than the average return rate for the total Aché diet,
which agrees with optimal foraging predictions (Hawkes, et al. 1982; Hill and Hawkes 1983).
The availability of shotguns does not cause certain low-ranked resources, such as small
monkeys and birds, to be included in the diet; rather, guns increase the return rates of all prey
species, so already low-ranked resources maintained their relative position (Hill and Hawkes
1983). Hill et al. (1987) take into account the desirability of specific nutrients, such as
protein, in the diet as opposed to just calories, which is often a criticism of prey choice
models (see above).
Winterhalder (1981) applies the prey choice model to the Cree foragers of northern
Ontario, who rely heavily on hunting for their subsistence. The Cree provide an interesting
test of the prey choice model because the resources that they exploit, such as moose and
caribou, waterfowl and fish, all require different technologies to procure. Therefore, the
hunting technologies (shotgun, 0.22 rifle, fishing nets) carried on a foraging trip determine
which resources are pursued, and the Cree often bring diverse weapons even if they set out to
hunt one species (Winterhalder 1981). Further, the introduction of technologies in the mid1900s such as snowmobiles and onboard motors for canoes have increased the distances and
34
prey types that the Cree seek to procure. Winterhalder’s (1981) study of the Cree provides
one of the best examples of how changing hunting and travel technologies impact forager
decisions. The introduction of motorized travel and guns within a short period brought about
a dramatic change, but prehistoric technological changes such as the appearance of
composite tools and projectile weapons, the bow and arrow, nets and snares, and horseback
riding would likewise have altered the efficiency with which humans exploited game.
One of the key components of the prey choice model is determining which species are
low- and high-ranked in an environment. Prey ranking typically is thought to be tied closely
to body size, with large taxa assigned a higher rank (see Hawkes, et al. 1982; Kelly 1995;
Simms 1987; Winterhalder 1981), as long as they are not unreasonably difficult to procure or
time-consuming to process (Byers and Ugan 2005; Jones 2004; Smith 1991). Prey ranking
among large game species can also be influenced by the age of animals in a population,
because adult animals tend to be larger and have a higher proportion of body fat (Broughton
2002; Munro 2004; Speth and Clark 2006; Stiner 1994). As a general rule, small-bodied
animals have typically been categorized as low-return because they provide a smaller
nutritional package than large game species (Broughton 1994a, 1994b, 1999). However, this
categorization of small-bodied prey as low-return has not traditionally taken into account
differences in handling costs.
Stiner and colleagues (Munro 2004; Stiner 2001; Stiner and Munro 2002; Stiner, et al.
1999; Stiner, et al. 2000) have shown that capture costs, and therefore return rates, vary a
great deal among small game animals, based on fundamental differences in their flight
behavior. This means that some small game could be much higher-ranked than others. For
example small, relatively sessile species (e.g., limpets and tortoises) have much lower
35
handling costs than small, fast-moving species (e.g. hares, birds and fish), even though their
caloric (and meat weight) values may be similar. Efficient exploitation of small, fast-moving
species require a considerable technological investment, such as snares or nets, and/or
cooperation of large groups of people (Cannon 2000; Jones 2006; Lupo and Schmitt 2002;
Madsen and Kirkman 1988; Madsen and Schmitt 1998; Schmitt, et al. 2004; but see Ugan
2005b for a discussion of small game in different habitats). Because of these differences in
search and handling costs, a small slow-moving animal has a higher return rate than a fastmoving animal of a similar size, and should be preferentially exploited by human hunters
(Stiner 2001; Stiner, et al. 2000). This is borne out by ethnographic data (Kelly 1995; Kuhn
and Stiner 2001: Table 5.1).
Some applications of prey choice models, and other optimal foraging models
discussed below, pertain to the question of resource depression, which occurs when prey
capture rates decline in response to harvesting pressures (Charnov, et al. 1976). Many
archaeological studies that employ prey choice models also address resource depression (e.g.
Broughton 1994a; Broughton 1994b, 1997, 1999, 2002; Butler 2000, 2001; Butler and
Campbell 2004; Cannon 2000; Grayson and Delpech 1998; Hill 2007; Jones 2004, 2006;
Manne and Bicho 2009; Munro 2004; Nagaoka 2001, 2002a, 2002b; Speth 2004; Speth and
Clark 2006; Stiner 2001, 2005, 2009; Stiner and Munro 2002, 2011; Stiner, et al. 1999;
Stiner, et al. 2000; Ugan 2005a). This is because resources at a site are fairly easily ranked
and change through time is assessed by the addition or removal of prey species in the diet.
Independent data on prey size diminution may also be used to check these interpretations.
Often, resource depression is linked to changes in local environments or climatic conditions,
but may also result from increased human population densities or resource intensification.
36
The archaeological examples cited above are far too numerous to discuss in detail in this
presentation. Suffice it to say, their application ranges from the Paleolithic through historic
periods on nearly all continents. The works listed above that relate more directly to
Paleolithic prey selection are discussed further in later chapters.
Patch Choice Model
The patch choice model is similar to the prey choice model, except it assumes that
resources are encountered in discrete patches, as opposed to evenly distributed across the
landscape. The model was developed by MacArthur and Pianka (1966) to determine which
patches an organism will utilize and has subsequently been applied to human foragers.
Patches are encountered by foragers randomly and sequentially on the landscape and are
ranked based on their return rates (Kelly 1995). Upon entering a patch, the forager's return
rate drops as resources are depleted. The patch choice model is often used in conjunction
with the marginal value theorem (MVT), which assumes that once the return rate from the
patch is less than the average rate of the surrounding environment (i.e., outside of the patch)
a forager will leave the patch and travel to the next one (Charnov 1976; Charnov, et al.
1976). The MVT adds travel time between patches and handling time within a patch, which
is not taken into account with patch choice alone. Application of the model requires more
information than the simple prey choice model described above.
Several ethnographic studies have used patch choice to understand recent human
foraging patterns (Beckerman 1983; Hawkes, et al. 1982; O'Connell and Hawkes 1984;
Smith 1991; Sosis 2002; Winterhalder 1981). The result from these studies are mixed, and
seem to suggest that foragers do not exclusively target the most profitable patches on the
37
landscape. Rather, patch exploitation varies on a daily or seasonal basis in response to
variations in patch profitability.
A benefit of the patch choice model is that patches can be defined quite broadly,
referring to the distribution of a single species (e.g., blackberry bushes), an ecosystem (e.g.,
marine, terrestrial, montane, deciduous forest), foraging strategy (e.g., fishing or hunting
large game) (Lupo 2007). Many archaeological applications of the patch choice model have
drawn on ethnographic studies that look at prey types that inhabit a specific ecosystem, for
example grassland or marine environments (Broughton 1999; Cannon and Meltzer 2008;
Jones 2007, 2009; Nagaoka 2002a, 2002b). These studies find changes in resource
exploitation as the result of increased “patchiness” of environments caused by climatic
change (Jones 2007, 2009), as well as shifts to more technologically difficult to exploit
environments, such as offshore areas (Nagaoka 2002a, 2002b), or the use of distant patches
as a result of resource stress (Broughton 1999).
Another archaeological application of the patch choice model seeks to understand
processing intensity of individual animal carcasses. The model is used in much the same way
as examining patches on a landscape, but the scale of analysis is different. Carcasses are
analyzed as individual patches and decisions are made about how intensively to process each
patch (carcass) before moving onto the next patch (kill) (Burger, et al. 2005; Nagaoka 2005).
Animal tissues vary in their nutritional return based both on their inherent caloric value and
the effort needed for extraction. In general, meat and organs have the highest returns,
followed by bone marrow, and finally bone grease (see values in Binford 1978; Kooyman
1990; Lupo 2006; Madrigal and Holt 2002 for a few examples). In exploiting the patch
(carcass), foragers initially should utilize the highest return parts, meat and organs, followed
38
by bone marrow, and only move on to processing labor-intensive bone grease (see Chapter 6
for a description of the process) if the processing costs are lower than the combined travel
and handling costs of seeking another prey item. Presumably, this occurs during times of
resource stress brought on by seasonal factors or long-term resource depression.
Burger et al. (2005) apply the MVT to predict foraging intensity across animal
carcasses. Using values from Binford (1978) and Madrigal and Holt (2002), the authors
construct food gain curves to predict the point at which certain low-return ungulate elements
should be processed for bone marrow. Burger et al. (2005:1151) note the nutritional
differences between lean protein and fatty bone marrow, which is easier to metabolize than
protein, and suggest that foragers will always seek some amount of marrow when processing
an animal. Foragers are expected to process marrow more intensively during times of
resource stress and when meat is most lean, though this can also vary depending on the
seasonal condition of an animal. The authors propose that a “stop element approach” can be
taken with archaeological assemblages, where different rates of marrow processing for lowutility elements are indicative changing butchery strategies, possibly in response to resource
stress (Burger, et al. 2005). The authors also note that increased rates of fragmentation may
indicate resource stress, either associated with marrow processing or bone grease rendering.
Obviously post-depositional factors that affect fragmentation would also have to be taken
into account in archaeological cases.
Nagaoka (2005) uses the patch choice model to examine increases in the processing
intensity of moa at the Shag River Mouth site in New Zealand, which dates to 1250-1450 CE.
The author uses the MVT to predict that moa should be processed more intensively when
resources were stressed in later time periods, which is well-established at the site (Nagaoka
39
2001, 2002a, 2002b). Following Wolverton (2002), she looks at the percent of unbroken moa
elements and the amount of bone marrow each element yields in order to determine if
elements were processed more intensively through time. Nagaoka (2005) determines that,
whereas large marrow-rich moa long bones were consistently broken throughout the
sequence, phalanges with small amounts of marrow are broken more frequently in later
layers. This trend indicates a rise in the intensity of carcass processing with time. The author
also looks for evidence of bone grease rendering by looking at changes in NISP:MNE ratios
(following Wolverton 2002). She finds no evidence for grease extraction, except on moa
tibiotarsi (Nagaoka 2005).
Many other archaeological studies examine carcass processing as a means of
intensifying animal resources, either in response to human or environmental pressures (e.g.,
Adouze 1987; Adouze and Enloe 1991; Binford 1993; Brink 1997; Broughton 1999; David
and Enloe 1993; Egeland and Byerly 2005; Hill 2007; Manne and Bicho 2009; Munro and
Bar-Oz 2005; Potter 1995; Stiner 2003b; Weniger 1987). Though these authors do not
necessarily present their results in terms of patch choice models, they follow the basic tenets
of patch choice and MVT, as they address changes in the frequencies of bone marrow or
grease processing through time in response to resource depletion.
Central Place Foraging Models
Central place foraging (CPF) models were developed by ecologists to understand
transport decisions by species that bring food resources to a home base, or central place, in
order to eat without interference, provision offspring, or store resources (Orians and Pearson
1979:156; Schoener 1979). A central goal of the models is to understand how resource
40
patches are chosen, how prey is selected, and how load size is determined (Orians and
Pearson 1979:156). The models predict that as distance from a central place increases, the
size of the load also increases in order to make travel costs worthwhile (Orians and Pearson
1979; Schoener 1979:913-914). This prediction seems to be valid for many non-human
predators (see Stephens and Krebs 1986:194-197).
CPF models were readily adopted by anthropologists, as movement of resources to a
home base is a typical human behavior. Anthropological applications of CPF models are
wide-ranging, from the transport of raw materials for stone tool production (Beck, et al.
2002), to the movement of plant resources (Barlow and Metcalfe 1996; Bettinger, et al. 1997;
Jones and Madsen 1989), to the transport of aquatic (Bird 1997; Bird and Bliege Bird 1997)
and terrestrial animal species (Bunn, et al. 1988; Egeland and Byerly 2005; Lupo 2006;
O'Connell, et al. 1988, 1990; O'Connell and Marshall 1989; Zeanah 1999, 2004). Each of
these studies attempts to assess the distance at which it becomes worthwhile for human
foragers to field process a given resource, and the amount of processing that yields optimal
economic gain. The formal model introduced by Metcalf and Barlow (1992:343) explores the
relationship between the round-trip travel costs of visiting a resource patch, the amount of
time spent procuring and field processing the resource, and the utility of the resource load.
The model predicts an inverse relationship between an increase in load utility from field
processing and the distance traveled before field processing becomes efficient (Metcalfe and
Barlow 1992:347). Therefore, field processing that results in a large increase in load utility is
more beneficial at shorter travel distances, and field processing resulting in small increases in
load utility only become worthwhile after longer distances are traveled. Metcalfe and
Barlow’s (1992) model can be most directly applied to simple, “structured” resources such as
41
seeds or mollusks that must be processed in a specific order (i.e., the removal of the outer
shell before the inner shell when processing some kinds of nuts). However, they recognize
challenges associated with utilizing more complex, “unstructured” resources such as animal
carcasses that can be processed in an almost infinite number of ways (e.g., after skinning, the
hind limbs, forelimbs, head parts or axial skeleton of an animal could be butchered next).
Cannon (2003) expands on Metcalfe and Barlow’s (1992) model by presenting a
“central place forager prey choice model” that takes into account differences of prey size,
transport distances, and processing costs. The author argues that though intensive field
processing may increase load utility, return rates are always higher for resources taken close
to home, regardless of processing costs (i.e., increased load efficiency does not fully
compensate for travel costs) (Cannon 2003:9). He also points out that while in general largebodied prey provides a higher return than small-bodied prey, there comes a point if large
game is locally unavailable that the search and transport time for such resources becomes so
costly that local small prey actually provides a higher return rate (Cannon 2003:12).
Applications of CPF models or those using similar rational have been fruitful in
explaining butchery and transport decisions that surround the processing and movement of
vertebrate remains from kill sites to a home base (Binford 1978; Bunn, et al. 1988; Egeland
and Byerly 2005; Lupo 2006; O'Connell, et al. 1988, 1990; O'Connell and Marshall 1989).
Of the three models, application of this one requires the most information a priori. Binford’s
(1978) groundbreaking ethnoarchaeological work with the Alaskan Nunamiut established a
baseline set of data that included utility indices for meat, marrow, and grease from different
skeletal parts of dall sheep (Ovis dalli) and caribou (Rangifer tarandus). The combined
values of the indices from the three kinds of tissue (meat, marrow, grease) constitute the
42
general utility index (GUI), which helps to predict the decisions the Nunamiut make
concerning the butchery, transport and storage of carcasses (Binford 1978:72). The GUI is
further derived into the modified general utility index (MGUI) that takes into account lowutility elements that may be transported along with high-utility portions (Binford 1978:74).
Based on the MGUI and frequency of various skeletal parts in an assemblage, Binford
(1978:81) introduced a set of utility curves that predict different body part transport
strategies. His study, which is discussed further in Chapter 6, was an important first step in
quantifying carcass transport decisions by human hunters. This study has influenced almost
all subsequent ethnographic and archaeological investigations that employ CPF models.
Many ethnographic studies of body part transport focus on the Hadza of Tanzania
(Bunn, et al. 1988; Lupo 2006; O'Connell, et al. 1988, 1990). Bunn et al. (1988) provide an
account of transport patterns, but do not track the relative utility of different body parts. They
find that appendicular elements tend to be transported more frequently to base camps than do
axial elements (Bunn, et al. 1988:450), and that much fluidity exists in defining locations as
“butchery sites” or “home base sites.” This is because Hadza hunters often process and
deposit bones at “snack sites” between the kill location and home base, frequently re-using
both kinds of sites for various activities (i.e., using old “home base sites” as “snack sites”)
(Bunn, et al. 1988:437). Thus, they caution archaeologists attempting to interpret transport
patterns that sites may have multiple usages over time. O’Connell et al. (1988; 1990) explain
Hadza transport strategies in terms of carcass size and which elements should be selected for
transport. Using a scalogram (Gutman matrix) analysis to construct a ranking system for
different body parts, they determine that though there is some variation among species,
vertebrae, scapulae, pelvises and upper limb bones tend to be preferentially transported from
43
kill sites to base camps (O'Connell, et al. 1988:138). In order to determine the number of
bones transported for each carcass, transport costs were calculated based on the average
weight of a carcass per human carrier, multiplied by the distance in minutes from the
butchering site to the home base (O'Connell, et al. 1988:132). In general, O’Connell et al.
(1990:311) find an inverse relationship between the number of elements moved and transport
costs. Additionally, they find that variation in transport exists between species of similar
sizes, particularly if the animals have different body forms (O'Connell, et al. 1990:310). The
significance of the studies done by O’Connell et al. (1988; 1990) are in dispelling ideas that
appendicular elements are typically transported preferentially over axial elements; rather,
there is variation depending on the prey species under consideration and prevailing
technology (e.g., the Hadza have cooking pots suitable for boiling). Further, they caution that
lumping different species with similar body sizes, while a common archaeological practice
due to identification constraints, can obscure species-specific transport decisions.
Lupo (2006) employs CPF models to evaluate how processing and transport costs of
certain anatomical portions of impala and zebra affect decisions made by Hadza foragers.
She measures utility using the food utility index, which is the total body part weight minus
the dried bone weight (after Metcalfe and Jones 1988). Lupo distinguishes between FUI(t),
the total food utility index equivalent to the FUI of Metcalfe and Jones (1988) and FUI(r), the
remnant animal products, calculated by subtracting the weight of the meat removed in the
initial filleting and the weight of marrow removed from FUI(t) (Lupo 2006:36). Lupo finds
that the kind of element (e.g., axial vs. appendicular) is less predictive of transport decisions
than FUI(r), and there are differences between impala and zebras. Hadza butchering
strategies for impala are fairly straightforward: foragers discard elements that can quickly be
44
depleted of meat and marrow (Lupo 2006:46). However, zebras tend to be processed only for
meat in the field, with the separated meat and bone subsequently transported to the
residential site, the latter for marrow and grease processing (Lupo 2006:47). Lupo (2006:48)
postulates that this is because of the specific structure of equid bone, which makes it difficult
to process in the field without heat. Her study refines observations made by O’Connell et al.
(1988; 1990) and Bunn et al. (1988) and provides a quantitative argument for understanding
different transport decisions made by the Hadza.
These ethnographic studies are increasingly sophisticated in their use of CPF models
or similar rationale. They provide insight into interpreting carcass transport decisions in the
archaeological record, while also cautioning against using modern examples as direct
analogs. The greater range of unknowns in archaeological cases must also be acknowledged.
The ethnoarchaeological studies also illustrate the range of carcass transport decisions made
by modern foragers depending on the prey species, number of participants in the hunt or
transport effort, distance between the kill and habitation sites, available technology, and local
climatic conditions. These ethnographic examples are instructive for archaeologists seeking
to evaluate different factors past foragers may have taken into account when making
decisions about carcass transport.
Following the ethnographic work discussed above, many recent archaeological
studies have adopted CPF models to evaluate body part transport decisions of past foragers
(Broughton 1999; Cannon 2003; Faith 2007; Marín Arroyo 2009; Nagaoka 2002b; 2005, see
also Speth 1991; Speth and Scott 1989 for discussions of the long-distance transport of large
game to horticulturalist sites). Broughton’s (1999) application of CPF models to the
Emeryville Shellmound faunas from the San Francisco Bay (2600-700 BP) set the stage for
45
many subsequent studies of large game body part transport. Average food utility index (FUI)
values for artiodactyls are established for each stratigraphic layer in order to determine the
economic utility of elements transported to the site. Relative skeletal abundance (RSA) is
calculated by dividing the number of identified specimens for each anatomical portion by the
number of times the part occurs in the body (Broughton 1999:58). Mean FUI values are
calculated by multiplying RSA and FUI values for anatomical portions (from Metcalfe and
Jones 1988) and dividing by the total RSA for the archaeological unit. Broughton (1999)
finds an initial decline in mean FUI of artiodactyl remains across the lower layers of
Emeryville Shellmound, then a subsequent increase in FUI in the later layers. The author
interprets this as lowered foraging returns following the depression of local artiodactyl
populations, and a decrease in body part selectivity as local resources were exploited more
intensively (Broughton 1999:59). Then, in the upper layers, fewer low-utility elements were
transported to the site, indicating that foragers traveled further from the site to procure large
game.
Nagaoka (2002b; 2005) applies Broughton’s (1999) methodology to forager decisions
surrounding the movement of moa and seal anatomical parts to the Shag River Mouth site.
She finds an overall increase in high-utility moa body parts through the sequence, indicating
moa hunting further from the site. A closer look at the data that separates out the highest and
lowest-utility elements indicates that there was a ubiquitous increase in high-utility elements,
and an initial increase followed by a decrease in the lowest-utility parts (Nagaoka 2002b,
2005). The author concludes that the lowest-utility elements were initially “riders”
transported to the site still connected to high utility elements, but as foragers began to process
carcasses more efficiently these elements were removed before transport in later periods. In
46
examining the transport of seal remains, Nagaoka (2002b) determines that in earlier periods
higher-utility elements are found at the site, and in later periods both low and high-utility
elements appear. She proposes the possible explanation that seals were hunted at distant
rookeries in later periods, but transport costs were different because of the use of canoes.
Faith (2007) expands on Broughton’s method by using it and the Shannon evenness
index (following Faith and Gordon 2007) to evaluate the representation of reindeer elements
at Grotte XVI, France during the Middle Paleolithic through Magdalenian. The author builds
on a study by Grayson and Delpech (2003; Grayson, et al. 2001), who noted an increase in
the frequency of reindeer in the later layers of Grotte XVI as a result of cooling summer
temperatures. Rather than investigating resource intensification, Faith (2007) tests the
hypothesis that foragers became less selective about reindeer body part transport in response
to increased abundance near Grotte XVI. He finds a decrease through time in the mean food
utility index of elements in the assemblage, in addition to an increase in skeletal evenness of
elements transported to the site. Faith’s (2007) analyses suggest that there was less selective
transport of reindeer parts to the site as prey availability increased in the region during the
late Upper Paleolithic (see Grayson and Delpech 2003; Grayson, et al. 2001).
Following the basic premise of Broughton (1999), Cannon (2003) examines changes
in the anatomical representation of ungulate remains from multiple sites in the Mimbres
Valley, New Mexico between about 400 and 1450 CE. The author calculates the mean FUI
for artiodactyl remains from different cultural phases similarly to Broughton (1999), then
compares the means of each phase though an analysis of variance (Cannon 2003). The author
finds that average FUI values are highest in the earliest and latest phases (Early Pithouse and
Classic Mimbres/Terminal Classic Mimbres) and are lower in the intervening periods,
47
indicating that low utility elements were selectively excluded at the beginning and end of the
sequence. It must be noted, however, that the sample sizes for these periods are extremely
small (n<16), a surprising contrast to the rigorous statistical tests applied. Cannon (2003)
attributes the initial decline in FUI values to increased hunting party size and the later
increase to foraging trips further from the analyzed sites.
Marín Arroyo (2009) examines central place foraging to understand site catchment in
the Cantabrian region of Spain during the Magdalenian (14,000-10,000 BP), with a focus on
red deer and ibex hunting. Following Cannon’s (2003) model, the author introduces time and
distance components to the model, determining that smaller-bodied ibex are an energetically
efficient hunting option if their habitat is 45% closer to a base camp than red deer habitats
(Marín Arroyo 2009:31). She also calculates that red deer are completely butchered on a 7.5
hour hunting trip, resulting in the transport of high-utility elements to the site. Finally, Marín
Arroyo (2009:31) determines that it is rarely efficient to process red deer bone marrow in the
field, unless it was consumed as an on-site snack. The author applies these assumptions and
topographic data to nineteen Magdalenian sites in Cantabria and finds a significant
relationship between topography and patch distance, and frequencies of highland- versus
lowland-dwelling species in the zooarchaeological assemblages (Marín Arroyo 2009).
The examples discussed above illustrate the potential utility of applying central place
foraging models to archaeological sites from diverse geographical areas and time periods.
The underlying assumptions allow for a quantitative evaluation of change in body part
transport strategies through an archaeological sequence. Obviously factors beyond travel
time can affect transport strategies (e.g., size of the foraging party, transport technologies
48
such as boats or vehicles), but establishing a trend or lack thereof is a first stage in evaluating
forager transport strategies.
Predictions of Hominin-Prey Relationships
Central to the use of models from evolutionary ecology that are outlined above is the
assumption that foragers seek to optimize their behavior if it will make them more
reproductively fit. Therefore, it follows that human foragers attempt to maximize the return
they get from their prey by exploiting the highest-ranked species or age groups available,
maximizing the gains from transport of resources depending on their distance from a site, and
through carcass processing. Substantial shifts in any of these areas of predatory behavior is of
interest, as it may be indicative of environmental pressures, resource depression or other
causes of socioeconomic change.
This dissertation seeks to apply the diet breadth or prey choice model, and the patch
choice and central place foraging models to the long sequence of zooarchaeological remains
from Klissoura Cave 1 in southern Greece. The prey choice model is applied to changing
proportions of low- and high-ranked prey (Chapter 5). The patch choice and central place
foraging models are employed for understanding changes in butchery patterns (i.e., marrow
processing) and body part transport decisions, respectively (Chapter 6). Data on
environmental change are used to qualify the interpretations of changes in prey use over
time. In addition to environmental impacts on the faunas available to foragers occupying
Klissoura Cave 1, evidence from elsewhere in the Mediterranean indicates that human
population densities were increasing through the Middle and Upper Paleolithic (Lahr and
Foley 2003), and into the Mesolithic. Such population growth would likely place increasing
49
pressure on the animal populations exploited by human hunters. In this context, it is predicted
that resource depression, if present, should be apparent in the dietary spectrum at Klissoura
Cave 1 over time. Expanding diets may be reflected by 1) an increase in low-ranked prey
types, including difficult-to-catch small animals such as hares and birds, as well as more
young ungulates, 2) an increase in high-utility ungulate body parts as hunters forage further
from the site, and 3) more intensive processing of carcasses, including opening long bones
for marrow and possibly even bone grease rendering. The following chapters explore these
lines of evidence to determine the extent to which the depression of food resources in
southern Greece resulted in changes in animal exploitation, and to what extent environmental
factors played a role in dietary change.
THE SAMPLE
The research presented in this dissertation is based on the vertebrate faunal
assemblages from Klissoura Cave 1, a stratified rock shelter near Prosymni in Peloponnese,
Greece. The site contains a large Middle Paleolithic component that spans Marine Oxygen
Isotope Stage (MIS) 5a at about 80,000 BP through the late Middle Paleolithic (ca. 53,600
cal BP), Early Upper Paleolithic (Uluzzian), Aurignacian, Epigravettian and Mesolithic, the
latter two periods being less well represented (Koumouzelis, Ginter, et al. 2001; Kuhn, et al.
2010; Stiner, et al. 2010). The long, relatively intact cultural sequence of Klissoura Cave 1 is
particularly important for diachronic comparisons of changes in hunting strategies and diet in
southern Greece during the late Pleistocene. The sample includes nearly 25,000 identified
remains from a long series of stratigraphically intact layers (mixed deposits were excluded
from the final analysis, see Chapter 2). To date, the analysis presents one of the largest, most
50
complete Paleolithic faunal sequences from Greece. Though one site cannot be taken as fully
representative of hominin behavior in an area, the data from Klissoura Cave 1 provide a
critical starting point for understanding changes in subsistence strategies in southern Greece
during the late Pleistocene. The robust sample provides evidence for dietary trends that relate
to local climatic fluctuations, as well as larger demographic changes that occurred as hominin
populations grew throughout Europe during the Paleolithic and later periods. The Klissoura
Cave 1 data are compared to published faunal data sets from contemporary faunal series in
Greece and the Mediterranean Basin to more fully understand late Pleistocene subsistence
strategies and dietary change.
DISSERTATION STRUCTURE
This dissertation is organized into eight chapters, beginning with an introduction to
the problem and the theoretical paradigm through which the data are interpreted. Chapter 2
provides an overview of Paleolithic research in Greece and a discussion of the excavations
(history and procedures) at Klissoura Cave 1. Background information about the
environmental conditions for the duration of the occupation of Klissoura Cave 1, with a look
at the Mediterranean region in general, and the local conditions in Greece specifically is
provided in Chapter 3.
Chapter 4 describes the analytical methods employed in this study and explores the
non-human taphonomic processes that may have affected the character of the faunas from
Klissoura Cave 1. A discussion of non-human taphonomic factors, including the potential of
in situ density-mediated attrition, is critical to understanding the formation of a faunal
assemblage and distinguishing trends in human faunal exploitation from natural processes.
51
Chapters 5 and 6 focus on human hunting patterns and bone modification at Klissoura Cave
1. Chapter 5 discusses prey choice and species abundance by looking at the frequency of
different taxa exploited at the site. Prey species are considered individually, as well as by the
categories of body size and flight pattern discussed above. Using diversity indices, changes
in species and prey group exploitation patterns are evaluated for trends. Alternative
indicators of changes in faunal exploitation patterns are presented in Chapter 6, focusing on
body part transport decisions, prey butchery patterns and age structure analysis. Chapters 5
and 6 draw heavily on expectations predicted by evolutionary ecology, using the prey choice,
central place foraging, and patch choice models to evaluate intensification efforts that
occurred at Klissoura Cave 1 during the late Pleistocene.
Chapter 7 examines inter-site variation of bone burning and fragmentation at
Klissoura Cave 1 to the extent that available excavation data permit. A series of intact hearth
structures, including dozens of clay-lined heart features and a possible habitation structure in
the Aurignacian layers, suggest important spatial variation in site use; this analysis will focus
on the richest, most intact layers. Finally, Chapter 8 offers a summary and synthesis of the
faunal results from Klissoura Cave 1, and a discussion of the overarching trends in faunal
exploitation patterns from the Middle Paleolithic through Mesolithic. The results from
Klissoura Cave 1 are then examined in the context of the archaeological record from Eastern
Europe and the greater Mediterranean region, with a special focus on resource intensification
in light of environmental variability and possible trends in human demography.
52
CHAPTER 2: KLISSOURA CAVE 1 AND
THE GREEK PALEOLITHIC
INTRODUCTION
Paleolithic research in Greece began rather late, if compared to parts of Western
Europe and Southwest Asia. This is often attributed to the historical interest in Classical
archaeology of the region, and the divide between classical and prehistoric studies in
universities (Runnels 1995b, 2001). Ultimately, this lack of attention early on could be
argued to be a positive development as many Greek Paleolithic sites have had the benefit of
being excavated using modern methods and technologies. Only a few small-scale excavations
or surveys were conducted prior to 1950, and unfortunately artifacts associated with these
early investigations were lost during the war years (Runnels 1995b, 2001).
Starting in the late 1950s, interest increased in the Greek Paleolithic. Surface finds in
the Argolid (NW Peloponnese) led to a series of unsuccessful cave excavations in the region
(e.g., Bialor and Jameson 1962). At the same time, Paleolithic surveys and excavations were
underway by a team in Thessaly (central Greece) (Milojčić, et al. 1965). The Middle
Pleistocene Petralona skull was subsequently discovered in the mid 1960s (Hennig, et al.
1981; Ikeya 1980; Kurtén and Poulianos 1977). The first comprehensive survey of northern
Greece was conducted in the early 1960s, which led to excavations of the early Middle
Paleolithic open-air site of Kokkinopilos, and the late Pleistocene rock shelters of
Asprochaliko and Kastritsa (Dakaris, et al.; Higgs and Vita-Finzi 1966; Higgs, et al. 1967).
Shortly thereafter, as part of a larger Neolithic and Bronze Age project on the Ionian islands,
Grava, an Upper Paleolithic rock shelter on Corfu, was excavated (Sordinas 1969). A salvage
53
operation was conducted at Kephalari Cave, a Middle and Upper Paleolithic site in the
Argolid, in the early 1970s. The first large-scale systematic excavation at a Greek Paleolithic
site also began at this time at Franchthi Cave, which spans the Upper Paleolithic through
Neolithic. The site was excavated from 1967 until 1979 and is located in the Argolid
(Jacobsen 1981; Jacobsen and Farrand 1987).
These projects ushered in the modern era of Greek Paleolithic archaeology, which has
witnessed a series of surveys and excavations primarily in Epirus (northwestern Greece) and
Peloponnese. Consequently, most of what we currently know about the Greek Middle and
Upper Paleolithic comes from northern Greece and the western half of Peloponnese (though
some "Lower Paleolithic," i.e. early Middle Paleolithic, tools were reportedly discovered on
Crete, see Strasser, et al. 2010).
Some early work on the Greek Mesolithic was conducted by Adalbert Markovits in
the 1920s, but his collections were thought to be missing until recently (Galanidou 2003).
The excavations at Franchthi Cave exposed an extensive Mesolithic occupation, including
multiple human burials and cremations (see below), which brought the Greek Mesolithic to
the attention of many prehistorians. Subsequently, a small handful of sites with Mesolithic
components were excavated and collections from old excavations are currently undergoing
analysis.
A perennial problem with Paleolithic archaeology in Greece is that the results of
many excavations are only partially, or never, published. The recent interest in Greek
prehistoric archaeology gained additional traction in the late 1980s, and has resulted in a
dramatic influx of information available to a much wider academic audience. New
excavations and analyses of previously excavated materials have yielded a growing body of
54
faunal and lithic data, in addition to the publication of some organic materials and human
skeletal remains.
In this chapter, I review the results of archaeological studies of the Paleolithic and
Mesolithic of Greece, focusing especially on recent work and mainly on cave sites. I will
emphasize the late Pleistocene and early Holocene, beginning at about MIS 5a (80,000 BP),
which marks the earliest occupations at Klissoura Cave 1. Open-air sites are excluded from
this presentation, unless they preserve faunal material or have radiometric dates (see
Ammerman, et al. 1999; Darlas 1999; Dousougli 1999; Papagianni 2000; Runnels 2001;
Runnels, et al. 2003; Runnels and van Andel 1993 for a discussion of some open-air Greek
Paleolithic sites). This chapter is organized geographically, starting with the sites in the
north, then moving to the Peloponnesian sites, and concluding with an in-depth description of
Klissoura Cave 1, the focus of this study. Sites included in the text appear in Figure 2.1; see
Figure 2.2 for ranges of dates for sites discussed below. All radiocarbon dates are calibrated.
NORTHERN GREECE
Asprochaliko
Asprochaliko, a Middle to Upper Paleolithic rock shelter in the Louros Valley, was
part of the early excavation efforts in northern Greece during the 1960s (Higgs and VitaFinzi 1966). Excavations occurred inside the rock shelter and on a slope outside of the
shelter, though the stratigraphy of the latter is questionable (Bailey, et al. 1983). The
sequence contains an Upper Paleolithic and two Middle Paleolithic components, and spans
about 100,000 to 26,000 BP based on thermoluminescence (TL) and radiocarbon dating
(Bailey, et al. 1983; Bailey, et al. 1992; Huxtable, et al. 1992).
55
Figure 2.1 Map of Greece with Paleolithic and Mesolithic sites mentioned in the text.
TL dates place the basal Mousterian (Middle Paleolithic) at Asprochaliko between
about 102,000 and 96,000 BP (Huxtable, et al. 1992). Tools in this older layer are “typical”
Mousterian and the assemblage is dominated by side scrapers and notches (Gowlett and
Carter 1997:445). The later Mousterian contains the same raw materials found in the older
56
layer but is technologically distinct (Papaconstantinou and Vassilopoulou 1997:461; Bailey
et al. 1983 calls it the "micro-mousterian"). Pseudo-Levallois points are found in the later
Mousterian layers. The technology used to manufacture such points is a combination of the
Levallois technique and a discoid method; the authors subsequently call it the Asprochaliko
method as it is not found elsewhere (Papaconstantinou and Vassilopoulou 1997:463).
“Asprochaliko flakes” manufactured by this method are considerably smaller than flakes
produced using the typical Levallois technique. The authors argue that it is difficult to place
the Greek Mousterian within a larger Eastern European chronology because of the variability
found in the region (Papaconstantinou and Vassilopoulou 1997:477).
The Upper Paleolithic lithics from Asprochaliko are presented by Bailey et al. (1983)
and Adam (1989; 1997). The Upper Paleolithic minimally dates to 26,350 +/- 1,900 BP, and
is defined as possibly Gravettian (Bailey, et al. 1983). The lithic sample is fairly small and
lacks evidence of production stages on site; no blades or blade cores are present. Bladelet
tools and endscrapers dominate the assemblage (Adam 1989, 1997). The authors note that
there is no Early Upper Paleolithic industry comparable to the Aurignacian or Uluzzian
(Bailey, et al. 1983:33), and a sterile layer of sediments indicates a hiatus between the Middle
and Upper Paleolithic components. A few worked bone specimens were recorded (Bailey, et
al. 1983).
The faunal assemblage contains 1,414 identifiable specimens, but only about forty
percent of these came from areas of known provenience (Bailey, et al. 1983:34). Despite this
qualification, the authors assert that the faunal composition changes through the sequence,
though it is always dominated by ungulates. The basal Mousterian contains abundant red
deer, and rhinoceros was recorded in the earliest levels (Bailey, et al. 1983:34). In the
57
Figure 2.2 Calibrated radiocarbon dates and ranges of dates for sites mentioned in the text. Cultural
periods are listed above their respective dates. Where radiocarbon dates are unavailable, placement of
cultural periods are estimated based on dated industries in Greece.
58
subsequent Mousterian, there is a fairly even representation of red deer and fallow deer. By
the Upper Paleolithic, red deer is still present, but ibex replaces fallow deer (Bailey, et al.
1983:34).
Theopetra Cave
Theopetra Cave in Thessaly contains a long prehistoric sequence, spanning the
Middle Paleolithic through the end of the Neolithic. The Middle Paleolithic layers were
initially thought to date to about 46,050 to 34,050 BP based on radiocarbon dating
(Facorellis, et al. 2001; Kyparissi-Apostolika 1999). Recent TL dates now place the Middle
Paleolithic layers at 124,000 +/- 16,000 and 129,000 +/- 13,000 (Valladas, et al. 2007:306).
This means that the earlier Middle Paleolithic deposits at Theopetra Cave are among the
oldest in Greece. Because they are much older than the Klissoura Cave 1 assemblages, they
are discussed only briefly here. Three Middle Paleolithic lithic phases were recorded at
Theopetra Cave (Panagopoulou 2000). The oldest (phase 1) is defined as a Quina industry,
followed by phase 2, which has Levallois blank production and a Mousterian toolkit. Lithic
phase 3 was initially defined as a terminal Middle Paleolithic or a Middle to Upper
Paleolithic transitional industry (Panagopoulou 2000), but recent dating indicates that the
these materials actually come from mixed deposits (Valladas, et al. 2007:307). A
stratigraphically higher TL date yielded an age of 59,000 +/- 6,000 BP, though little cultural
material was noted in this layer (Valladas, et al. 2007).
The stratigraphy of Theopetra Cave is difficult to understand as it includes a series of
hiatuses and cryogenic events (Karkanas 2001). The recently published TL dates provided by
Valladas et al. (2007) will undoubtedly lead to a reinterpretation of the stratigraphy. Many
59
long hiatuses are apparent at the site, most of which relate to the early Middle Paleolithic
layers. Early Upper Paleolithic deposits are mixed with Middle Paleolithic layers (Adam
1999a). There is a hiatus in the sequence during the last glacial maximum, and the
subsequent Upper Paleolithic/Epipaleolithic deposits range between 17,850 +/- 700 and
13,000 +/- 200 BP (Kyparissi-Apostolika 1999). The Mesolithic layers are largely intact and
date to between 11,400 +/- 1,250 and 8,815 +/- 215 BP (Kyparissi-Apostolika 1999).
Lithic raw materials during the Upper Paleolithic and Mesolithic layers are primarily
radiolarite, though some non-local flints were also noted in the Upper Paleolithic (Adam
1999a). The Upper Paleolithic industries contain a low ratio of tools to debitage, and few
cores (Adam 1999a). The most common tool types are unilaterally backed bladelets,
retouched bladelets, and only a few blades, endscrapers and truncations are present (Adam
1999a:267). The author suggests based on the amount of debitage that many of the tools were
made at the site. In the Mesolithic layers there is no evidence for the production of laminar
debitage (Adam 1999a:267). Tools include truncations, notches and flakes with alternate
retouch. In general the results on the Upper Paleolithic and Mesolithic industries remain
preliminary.
To date, slightly fewer than 4,500 faunal remains from the Paleolithic and Mesolithic
layers at Theopetra Cave have been analyzed and published, of which 436 are identifiable
(Newton 1999). The author notes high incidences of post-depositional breakage of the
materials, with only a few green (fresh) bone breaks (Newton 1999:116). Human
modifications on the bones include some burning damage and a few cut marks, and rare
carnivore damage is also present. A range of ungulate species is recorded, including ibex,
equids, red deer, fallow deer, and roe deer. Ibex and equids are only found in the Paleolithic
60
layers (Newton 1999). Carnivores comprise about a fifth of the assemblage and include large
and small forms such as bear, hyena, wild cat, wolf, fox, badger, and marten. Small mammals
such as hare and beaver are found at Theopetra Cave, as are avian taxa including corvids,
rock dove, partridge, a few aquatic birds, and some remains of small owls and diurnal raptors
(Newton 1999). A generalized “reptile” category occurs mainly in the oldest Paleolithic
layers, but it is unclear which species are included and how they may relate to human
activities in the cave. Newton (1999) attempts to define trends through four major
stratigraphic sequences, but the samples are rather small and prohibit definitive conclusions.
Two human skeletons were discovered at Theopetra Cave, one from the late Upper
Paleolithic, which yielded a date of 16,500 +/- 500 BP on bone, and the other from the
Mesolithic layers within a pit that contains charcoal dating to 10,420 +/- 180 BP (the
skeleton itself was dated to 8,975 +/- 325 BP) (Stravopodi, et al. 1999:270). The late Upper
Paleolithic human remains were disturbed at some point in the past and only a few long bone
elements and the calvarium were recovered; they are thought to have belonged to an adult
male. The Mesolithic human remains are almost completely intact (Stravopodi, et al. 1999).
These remains are from a young woman between the ages of 18 and 20, who suffered from
slight nutritional deficiencies, possibly a lack of iron (Stravopodi, et al. 1999).
A final notable feature from Theopetra Cave is a series of four left footprints,
probably from two children between the ages of two and four years old (Manolis, et al.
2000). The prints are from the Middle Paleolithic layers that date in excess of 120,000 BP
(Valladas, et al. 2007). Based on experimental studies using stereophotography and
photogrammetry, the authors conclude that two of the prints were probably made by an
individual wearing some kind of foot covering, while the third was not. The fourth print is
61
somewhat obscured and might indicate an overlap of prints from the two different individuals
(Manolis, et al. 2000).
Kastritsa
Kastritsa is an Upper Paleolithic rock shelter that was discovered and excavated as
part of the work conducted in northern Greece during the 1960s (Higgs, et al. 1967). Five
stratigraphic layers were uncovered spanning 23,930-19,000 BP (AMS dates on charcoal)
(Galanidou and Tzedakis 2001; Galanidou, et al. 2000), though most of the archaeological
materials are in the upper three layers that post-date 22,280 BP. The lithic industries were not
assigned to a specific cultural period, though they encompass a range defined elsewhere as
Gravettian (Adam 2007). Most of the tools in the lowest layer (Stratum 9), which is a beach
deposit, are endscrapers; there are few bladelet tools and no evidence of core exploitation
(Adam 1997). Laminar blanks were produced in all layers. This period probably represents
brief visits to the site. Starting in the overlying archaeological layer (Stratum 7), and
especially in Stratum 5 the shelter was used much more intensively. Shouldered points and
backed bladelets that were hafted appear at this time (Adam 1997). Stratum 5 contains stone
lined hearths and a stone tool industry dominated by flakes; microgravettes are also present
(Galanidou 1997, 1999). A small fraction of the raw materials in this layer were transported
up to 50 km from the Voidomatis River area (Adam 2007). Refitting studies of the lithics
indicate no preserved spatial variation in site use in Stratum 5, rather it is interpreted as a
“horizontal palimpsest” of multiple occupations in a short time period (Galanidou 1997,
1999). Stratum 3 contains more exotic raw materials, also from the Voidomatis River (Adam
2007). The technologies in this layer are slightly different; there is an increase in cores and
62
the microburin technique appears. Backed bladelets are common (Adam 2007). In the
uppermost layer, the microburin technique persists (Adam 1997). Overall the stone tool types
found at Kastritsa are especially diverse, even as compared to other sites in Epirus and those
in Peloponnese (Elefanti 2003).
The Kastritsa assemblages include an abundance of worked bone and antler, as well
as ornaments, in Stratum 5 and above (Adam and Kotjabopoulou 1997; Bailey, et al. 1983;
Kotjabopoulou and Adam 2004). Antler points are more common than bone points (Bailey,
et al. 1983). Seven perforated red deer canines were recovered, one of which is decorated
with incised grooves, in addition to three grooved long bone fragments (Kotjabopoulou and
Adam 2004). Despite the fact that Kastritsa is over 60 km from the coast, about two dozen
perforated Cyclope and Dentalium shells were found. Finally, two perforated serpentinite
“beads” were excavated in Stratum 5 (Kotjabopoulou and Adam 2004).
The faunas of Kastritsa were extensively studied by Kotjabopoulou (2001). A total of
4,623 specimens were identified to body class or species, most of which come from the upper
three stratigraphic layers. The assemblages are dominated by red deer, followed by birds,
wild ass, caprines, and carnivores. Wild pig, roe deer and lagomorphs were also recorded in
small numbers (Kotjabopoulou 2001). Water birds (family Anatidae, which includes ducks,
geese and swans) occur in the assemblages, though their proportions are unclear (Bailey, et
al. 1983; Kotjabopoulou 2001). No temporal trends related to species representation are
apparent at Kastritsa. High-utility bovid and cervid body parts are the most common at the
site, indicating the preferential transport of these elements (Kotjabopoulou 2001:77).
Conversely, equid anatomical profiles indicate a dominance of heads and hooves. This is
interpreted as either being the result of scavenging or of butchering at the site and moving
63
higher utility parts elsewhere (Kotjabopoulou 2001:77). However, the author notes that
diagenesis may have affected the assemblage (Kotjabopoulou 2001:92) which may explain
some relationships between utility indices and survivorship. Using data from cervid tooth
eruption, Kotjabopoulou (2001:88) determines that the site was occupied during warm
months.
Klithi
Klithi, in Epirus, is one of the most intensively studied and thoroughly published
Paleolithic sites in Greece (e.g., Adam 1989; Bailey 1997, volumes 1 and 2; Bailey 1999;
Elefanti 2003; Galanidou 1999). Twenty-two AMS dates on bone collagen, charcoal and
burned bone range from 20,250 +/- 1,200 to 16,400 +/- 550 BP, though most dates fall
between 19,700 and 16,225 BP (Bailey and Woodward 1997; Gowlett and Carter 1997).
Spatial and temporal resolution at Klithi is low, partly because isolated trenches were
excavated in multiple areas across the site, and because stratigraphic layers could not be
correlated between the trenches (Wenban-Smith 1997). Consequently, analyses of the
artifacts treat all layers of the site as a single unit.
Local, low-quality raw materials dominate the lithic assemblages at Klithi (Roubet
1997). Backed bladelets are the most common tool type at the site (Adam 1997, 1999b;
Elefanti 2003; Roubet 1997, 1999; Roubet and Lenoir 1997). The microburin technique is
found throughout the site, in addition to the “transversal Klithian fracture (TKF)” which is an
alternate method of removing the ends of backed bladelets (Roubet 1999; Roubet and Lenoir
1997:157-161). Endscrapers are also common, in addition to notches and denticulates,
piercers and burins, side scrapers, and composite tools (Elefanti 2003; Roubet 1997). Use-
64
wear studies indicate that some backed bladelets were used as barbs on projectiles, for
working hides, or cutting/scraping bone or wood (Moss 1997).
A large number of organic artifacts (osseous tools and debitage) were recovered at
Klithi (Adam and Kotjabopoulou 1997; Kotjabopoulou and Adam 2004). Needles are
common (n = 21) as are awls (n = 20). Spatulas and bone points were also recorded (Adam
and Kotjabopoulou 1997:246; Kotjabopoulou and Adam 2004). The analysts recorded two
antler tools and eight perforated red deer canines, which is notable because only eleven
cervid specimens were recorded in the faunal assemblage (Adam and Kotjabopoulou 1997;
Gamble 1997; Gamble 1999). Nearly two hundred marine gastropods were recovered, most
of which are perforated. Cyclope is the dominant species, followed by Homalopoma
sanguineum, and a few occurrences of other species including Dentalium (Adam and
Kotjabopoulou 1997:251; Kotjabopoulou and Adam 2004:40).
Over 140,000 faunal specimens were identified at Klithi. The remains are described
as well-preserved but highly fragmented (Gamble 1997:210). Gamble (1997; 1999) notes the
extreme dominance of ibex and chamois at the site; these species have an NISP count over
12,000, the next closest taxon or class-specific identification is “indeterminate rodent” (NISP
= 316), followed by birds (NISP = 202). Indeed, occupants of Klithi focused on ibex hunting
in the classic sense described by Straus (1987) (Gamble 1997; Gamble 1999; PhocaCosmetatou 2003a). Analysis of anatomical representation indicates a bias against low-utility
parts (Gamble 1997; Gamble 1999). Butchery damage is common in the assemblage;
occurring on 13.4% of the remains (Gamble 1997:219). Age profiles based on bone fusion
data suggest that the majority of ibex in the death assemblage are adults (4-8 years), whereas
tooth eruption and wear data indicate that juvenile and young adult (0-4 years) and old (8+
65
years) groups are the most well-represented. The author proposes that the hunters at Klithi
followed a gourmet transport curve when prime-age adults were hunted, and a bulk utility
curve when old or young individuals were killed (following Binford 1978). The author does
not, however, address the effects attritional processes may have had on the assemblage.
Based on the occurrence of fetal or neonate remains and bone fusion patterns, the site was
probably occupied in spring, summer and autumn (Gamble 1997).
Boïla
Boïla rockshelter is a late Upper Paleolithic site in the Voidomatis Gorge. The
deposits are fairly shallow and four archaeological layers or sub-layers were noted. Unit IV is
undated and contains Epipaleolithic or Mesolithic tool types. All other layers contain stone
tools similar to the Balkan Late Epigravettian (Kotjabopoulou, et al. 1999:202). Unit IIIb
ranges between 13,000 +/- 1,350 and 11,850 +/- 600 BP based on dated charcoal and burned
bone, respectively. Unit IIIa yielded a date of 14,800 +/- 750 BP and Unit II 16,550 +/- 600
BP, both on burned bone (Kotjabopoulou, et al. 1997:429). These dates should probably be
regarded with some caution, as dates on burned bone are often too young, though the dated
carbon in Unit IIIb is fairly close to the date on burned bone in that layer, and the stone tool
industries match well with what is expected during this time period in Greece.
Diachronic change is apparent in the stone tool industries at Boïla. In the lower layers
(Units II-IIIb), cores are larger and there are more unilaterally backed bladelets. In Unit IV,
the microburin technique appears, along with Sauveterre points and microgravettes, and core
size is much smaller, suggesting a more intensive use of raw materials (Kotjabopoulou, et al.
1999:206). In all layers the lithic assemblages indicate that the site was primarily used as a
66
hunting camp where tool maintenance occurred (Kotjabopoulou, et al. 1999:206). Units IIIIIb at Boïla correlate technologically with Stratum 1 at Kastritsa and lithic phases IV-VI at
Franchthi Cave (see below).
A little over fifty ornaments were recovered from Boïla (Kotjabopoulou and Adam
2004). These were made from marine and freshwater mollusk shells, most of which are
perforated Cyclope. Only one modified red deer canine was recorded, and one steatite bead
was recovered (Kotjabopoulou and Adam 2004).
The faunal assemblage at Boïla is small and highly fragmented, with 1,203 identified
bones and teeth (Kotjabopoulou 2001:223). Ungulates dominate the remains, specifically
ibex and chamois, followed by red deer. Birds, fish, lagomorphs, and carnivores are also
present in the assemblage in lower frequencies (Kotjabopoulou 2001:223). According to
Kotjabopoulou (2001) humans at Boïla either had similar body part transport strategies for
cervids or caprines, or caprines were hunted at a distance from the site and cervids were
taken close to the shelter (depending on if unidentified ungulates are added to the
calculations) (Kotjabopoulou 2001:123-124). The analyst offers no analysis of taphonomy or
potential attritional processes affecting the faunas. The lack of this information combined
with the small sample sizes indicates that her conclusions should be viewed with some
caution. Young ungulates are common in the assemblage, and the faunal remains indicate
that the site was probably used between late spring and mid-summer (Kotjabopoulou 2001).
Grava
Grava is an undated rockshelter on the southern part of Corfu that yielded
diagnostically Upper Paleolithic stone tools (Sordinas 1969). Corfu was connected to the
67
mainland until after the last glacial maximum (Van Andel and Shackleton 1982), so
Paleolithic populations could have moved freely onto the “island” without crossing water.
The stone tools have recently been re-analyzed by Adam (2007) who found many similarities
to the industries from Strata 3 and 1 at Kastritsa, which coincide temporally with the
Gravettian. In general, blades are more common than bladelets in the assemblage. Tool types
include unilaterally backed bladelets, endscrapers made from retouched blades, and burins.
The author notes that both the tools and the technological styles are quite similar to those
observed at Kastritsa (Adam 2007:155-156). Worked bone was also recovered at Grava,
which includes a bone point and an incised bone (Sordinas 1969). A perforated red deer
canine was also found, along with one ochre-stained pebble (Adam 2007).
The faunal assemblage was not studied in depth, though Sordinas (1969) notes that it
was abundant. Aurochs are common, and other ungulate species include red deer, fallow
deer, roe deer, variously-sized equids, and wild pig. Snails and small mammals such as hare
and rodents were recovered, as well as fox, hyena, badger, and mustelids. Sordinas
(1969:399) notes “the great number of large birds which (he) was not able to identify.” Based
on other Paleolithic Greek faunas, I suspect that at least some of these remains were from
great bustard (Otis tarda).
Sarakenos Cave
In central Greece, on the shores of the now dried Lake Kopais, Sarakenos Cave
preserves a sequence that spans the Paleolithic through the Late Helladic period (Sampson, et
al. 2009). Though the authors discuss the Paleolithic and Mesolithic periods, they mostly
focus on the rich Neolithic layers. The Paleolithic occupations of the site were fairly
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ephemeral, and few faunal remains were recovered (Sampson, et al. 2009:228). Some
potential Early Upper Paleolithic or Middle-to-Upper Paleolithic transitional tools were
recovered from an undated layer of Sarakenos Cave. These include Levallois points and
flakes, and the authors compare them to transitional industries from Temnata Cave in
Bulgaria (Sampson, et al. 2009:214). Sampson et al. (2009) also note side scrapers,
denticulated-notched pieces and a blade thinned by the Kostenki technique, but the authors
do not provide measurements or further comparisons in support of these conclusions. Late
Paleolithic stone tools are tentatively assigned to the Epigravettian period, and include a
retouched truncation made from an obsidian blade (Sampson, et al. 2009:214). A radiocarbon
date on charcoal from this layer dates to 14,750 +/- 700 BP.
Numerous hearth features were uncovered in the Lower Mesolithic layers, which
yielded a tight series of radiocarbon dates ranging from 10,315 to 10,435 BP (Sampson, et al.
2009:213-214). The Late Mesolithic deposit dates to between 8,895 and 9,020 BP (Sampson,
et al. 2009:214). Sarakenos Cave was more intensively occupied during the Lower
Mesolithic, less during the Late Mesolithic (Sampson, et al. 2009:228). Further, during the
Late Mesolithic, occupants used local raw materials for the manufacture of stone tools, which
Sampson et al. (2009:228) suggests was a result of local groups becoming isolated. The
authors note that at this time big game hunting was replaced by the exploitation of birds and
plant foods, but they provide no quantitative evidence for this assertion.
Sidari
Sidari on Corfu contains Mesolithic and later horizons. Discovered in the 1960 the
Mesolithic component of the site is a shell midden with a radiocarbon date of 8,750 +/- 800
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BP (Sordinas 1969, 2003). Lithics from the site are mostly of flint; the source of the most
common type is unknown, though the author postulates that perhaps they are nonlocal
(Sordinas 2003:91). Non-geometric microlith technology is common, probably destined for
use in composite tools, and retouched pieces mostly include points, though scrapers, burins,
and other tools occur in low numbers. Backed bladelets are almost entirely absent from the
assemblage (Sordinas 1969, 2003). The author notes that the microlithis from Sidari are
unlike those found on the Greek mainland. Sordinas (1969:405) notes that small game is
present in Mesolithic level of the site, but the majority of the animal remains are from
Cardium edule (cockle shells). No quantitative values are provided for the shells or small
game.
Cave of Cyclope
The Cave of Cyclope is located on the southwestern part of the island of Youra in the
Aegean. The site contains rich Neolithic and Mesolithic deposits (Sampson 1998). The
Lower and Late Mesolithic layers preserve abundant faunal remains, hearth features, land
snails, marine shells, fishhooks and lithic materials, as well as a human skull from an older
female in the lowest stratigraphic layer (Sampson 1998). Three radiocarbon dates from the
Cave of Cyclope place the Lower Mesolithic occupation between 10,335 and 10,530 BP,
while dates from the Late Mesolithic range from 9,080 to 9,810 BP (Sampson, et al.
2003:125).
The Mesolithic lithic assemblage from the Cave of Cyclope is fairly small (n = 179).
Raw material types include nine different kinds of flint, chalcedony, quartz, and a few pieces
of obsidian (Sampson, et al. 2003). Cores are found in the assemblage, as well as
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endscrapers, retouched flakes and notched tools; a burin and two backed pieces were also
recorded (Sampson, et al. 2003:127). Evidence of tool manufacture at the site is apparent for
all raw material types except obsidian. The obsidian was chemically sourced to the island of
Melos, which is also the source of obsidian found in the Mesolithic layers of Franchthi Cave
in the Argolid (see below). Obsidian artifacts were not manufactured at the Cave of Cyclope;
pieces include eight tools, four blades, two flakes, and only one chip (Sampson, et al.
2003:127). The authors note that tools made of obsidian are different forms than those made
of other materials, and include crescents, a backed blade, a truncation, and a trapeze
(Sampson, et al. 2003:127).
A large osseous tool assemblage was recovered from the site (n = 78 from all
Mesolithic layers) and includes fish hooks, bi-points (or gorges), and bone points (MoundreaAgrafioti 2003). Most of the tools were recovered in Late Mesolithic layers. Bi-points, or
gorges, are the most frequent tool type in the Mesolithic layers. The author compares them to
Paleolithic points: splinters that taper toward one or both ends, constructed of small or large
animal long bones or antler (Moundrea-Agrafioti 2003:138). Bi-points are thought to have
been used for fishing, along with hooks, meaning that there are two classes of bone fishing
artifacts found at the site. Hooks are the second most common bone implement at the Cave of
Cyclope and range between 1-4 cm, though two are 5-6 cm long. Some of the hooks are
broken but many are intact (Moundrea-Agrafioti 2003). Fish hooks were constructed mainly
from ungulate long bone diaphyses, or the bones of small animals or birds. One is made of
deer antler, a species that probably was not found on the island (Moundrea-Agrafioti
2003:134).
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An abundance of faunal remains, including mammals (NISP = 13,350), birds (NISP >
850), and especially fish (NISP > 17,000), were recovered from the Mesolithic layers at the
Cave of Cyclope. The faunas in the Lower and Late Mesolithic layers differ from one
another. Caprines, mostly goat but also some sheep, dominate the mammalian assemblage of
both of the Mesolithic occupations, and suids are present in small amounts in the Lower
Mesolithic. Based on biometric and mortality data, Trantalidou (2003:170) postulates that
goats in the Lower Mesolithic were in a stage of proto-domestication, while those in the Late
Mesolithic were domesticates brought from the mainland. Land snails are also common in
the Lower Mesolithic layers, but the author does not consider their dietary contributions.
Avian remains include Cory’s shearwater, Manx shearwater, cormorant, shag, sparrow hawk,
quail, great bustard, turtle dove, little owl, and raven (Trantalidou 2003:159). The author only
discusses Manx shearwater and great bustard remains in detail, though their abundances
relative to the other faunal remains are not discussed. It is unclear if their frequencies
changed through the sequence.
Trantalidou (2003:144) suggests that fish dominate the Lower Mesolithic, but
published data on the fish remains (Mylona 2003; Powell 2003) indicate that fish are much
more common in the Late Mesolithic. The cranial and vertebral remains of fish were studied
separately, the former presented by Powell (2003) and the latter by Mylona (2003). The
conclusions of the two studies vary slightly, but their overall findings are the same. At least
29 species of fish were exploited at the Cave of Cyclope; the most common families include
sea bream, groupers, mackerels and tuna, and mullets (Mylona 2003:184; Powell 2003:173).
Both studies indicate that a wider range of species were identified in the Late Mesolithic
72
layers (Mylona 2003; Powell 2003), which may or may not be the result of larger sample
sizes in these levels.
SOUTHERN GREECE
Recent work on the Mani Peninsula in southern Peloponnese has yielded several
Paleolithic cave sites. Some of these sites are in advanced phases of excavation and
publication (Kalamakia, Lakonis, Apidima) and can be discussed at some length below.
Other sites on the Mani Peninsula have only undergone survey or initial testing and therefore
cannot be covered in detail here (see Darlas and Psathi 2008 for some recently discovered
Upper Paleolithic sites in the region).
Kalamakia Cave
Kalamakia Cave is a karstic limestone cave on the coast of southern Peloponnese that
preserves Middle Paleolithic deposits. Seven stratigraphic layers were excavated and
archaeological materials are found in layers III, IV and V (layers with lower numbers are
stratigraphically older) (Darlas and de Lumley 1999, 2004). The excavators believe that the
lowest layer was deposited in MIS 5e, despite a marine shell in layer II that was dated to
109,000 +/- 14,000 BP using the uranium disequilibrium method (Darlas and de Lumley
1999, 2004). The earliest archaeological evidence probably dates to MIS 5c, and a
radiocarbon date from layer IV provides a minimum age of 40,050 years (Darlas 2007).
Initial studies of the Kalamakia lithics indicated that there was a slight difference
between the lower and upper archaeological layers. The artifacts from the lower levels were
defined as typical Mousterian, with frequent high-quality Levallois flakes and numerous
73
flakes with facetted striking platforms (Darlas and de Lumley 1999). Conversely, artifacts
from the upper level belong to a more “Mousteroid” industry because they are of moderate
quality and do not belong to a standard Mousterian type. There is also a lack of facetted
platforms and low frequency of Levallois flakes (Darlas and de Lumley 1999). Subsequent
studies of the lithics have stressed that tools in the assemblage are typical Mousterian made
with Levallois debitage (Darlas 2007; Darlas and de Lumley 2004). Raw materials include
flint from 12-20 km from the site, and andesite, quartz and quartzite (Darlas 2007). Despite
the abundance of Levallois debitage in the assemblage, few points were recovered (Darlas
2007; Darlas and de Lumley 2004), which may indicate that the site was used for tool
manufacture.
A unique feature of the Kalamakia Middle Paleolithic assemblage is the presence of a
few shells of Callista chione (a smooth-shelled large clam). The shells were probably not
consumed, rather they were retouched and possibly used as scrapers (Darlas 2007; Darlas and
de Lumley 2004). This phenomenon is also reported at Grotta dei Moscerini in Italy (Stiner
1994:187-188 and references therein).
Mammalian faunal remains are in the process of being analyzed and published. A
wide range of ungulates are recorded at Kalamakia, dominated by wild goat, chamois and
fallow deer, but also including extinct elephant and rhinoceros, wild pig, red deer, roe deer,
and a large bovid (possibly aurochs or bison) (Darlas 2007; Darlas and de Lumley 1999;
Gardeisen, et al. 1999). At this point no trends are apparent in the faunas (Darlas 2007).
Ample carnivore remains in some of the layers indicate that the site alternated between a
human occupation site and carnivore den (Darlas and de Lumley 2004). Large and small
carnivores are found at the site, including brown bear, leopard, wolf, Eurasian lynx, wild cat,
74
fox and mustelids (Darlas 2007; Darlas and de Lumley 1999, 2004; Gardeisen, et al. 1999).
Microfauna are abundant in the Kalamakia stratigraphic series and include 59 species of
small mammal, amphibians, reptiles, and moderate amounts of tortoise. Much of the
microfauna is thought to be refuse from raptors or owls that nested in the site (Roger and
Darlas 2008a). A range of avian fauna was recorded in layers IV and V. Avian NISP counts
are low (NISP = 95) but represent twenty-one species (Roger and Darlas 2008b). The most
common species in the assemblage is quail, followed by rock partridge and pigeon, though
several species of diurnal raptor and owl were also recorded (Roger and Darlas 2008b). No
human modifications, such as cut marks or burning damage, are apparent on the bird remains
and only one incidence of raptor digestion was recorded. The authors suggest that the most
widely represented avian taxa, such as the partridges, quails and pigeons (which also appear
at other Greek Paleolithic sites) were collected by humans, while smaller birds represent the
remains of raptor meals (Roger and Darlas 2008b).
Hominin remains were identified in layer IV and include several Neandertal teeth, a
cranial fragment, fibula, and lumbar vertebrae (Darlas 2007).
Lakonis I
Lakonis I, a limestone cave in the southern Peloponnese, is reported to contain
stratified Middle (Units IV to Ib) through “Initial” Upper Paleolithic (Unit Ia) deposits. TL
and U-series dating of beach rock and speleothems indicates that the earliest Middle
Paleolithic was deposited between 130,000 and 120,000 BP (Elefanti, et al. 2008;
Panagopoulou, et al. 2002-2004). A series of AMS dates on charcoal place the late Middle
Paleolithic and earliest Upper Paleolithic between 46,600 +/- 6,050 and 38,600 +/- 2,450 BP,
75
though the authors note that the range of dates are statistically indistinguishable between the
two layers (Elefanti, et al. 2008; Panagopoulou, et al. 2002-2004).
Raw materials at Lakonis I are mostly local, found within about 5-10 km of the site
(Panagopoulou, et al. 2002-2004:331). The Middle Paleolithic layers are defined by the
production of blanks using the Levallois laminar method as well as non-Levallois methods.
The most common tool types are side scrapers and retouched Levallois and Mousterian
points; bifacial scrapers, knives and points are also found (Panagopoulou, et al. 20022004:336). Most points were produced using châpeau de gendarme platform preparation
(Panagopoulou, et al. 2002-2004:332).
The uppermost archaeological deposits (Unit Ia) contain a mix of Middle and Upper
Paleolithic tool types and technological styles (Elefanti, et al. 2008; Panagopoulou, et al.
2002-2004), which may indicate some mixing in certain parts of the site. However,
articulated faunal remains, refitted lithics, and micromorphology are taken to indicate that the
layer is stratigraphically sound (Elefanti, et al. 2008; Panagopoulou, et al. 2002-2004). Raw
materials in the Initial Upper Paleolithic layers are mostly local, and flint is much more
common than in underlying deposits (Elefanti, et al. 2008:89). Levallois and prismatic blank
production techniques are found in the layer. Bladelet manufacture is common, using two
different chaînes opératoires (Elefanti, et al. 2008:92). Stone tool types include Mousterian
points and notches/denticulates, which are considered Middle Paleolithic, though
manufactured as part of prismatic core production, similar to the Central European
Bohunician (Elefanti, et al. 2008:92). The most common tools in Unit Ia are Upper
Paleolithic in nature: retouched flakes, blades and bladelets, as well as burins, endscrapers,
and a few Aurignacian scrapers and blades (Elefanti, et al. 2008:92). Panagopoulou et al.
76
(2002-2004) argue that the IUP industry is indicative of an in situ transition from Mousterian
to Upper Paleolithic technologies.
The faunal remains from Lakonis I are still undergoing analysis, and preliminary
publications stress the extreme fragmentation of the assemblage (Panagopoulou, et al. 20022004). Panagopoulou et al. (2002-2004:339) note that as of 2001, 4.4 kg of faunal remains
were recovered from Lakonis I. Of this, 818 specimens are bone or dental specimens larger
than 2 cm. In this subset 73 specimens were identified (Panagopoulou, et al. 2002-2004:339).
The remains exhibit a high frequency of burning damage, with 67% (NISP = 31) of the
identifiable remains burned (Panagopoulou, et al. 2002-2004:340). The small size of the
assemblage makes stratigraphic divisions impossible, though the authors do attempt to
discuss anatomical representation. Identified taxa include fallow deer, red deer, roe deer and
pig, as well possibly aurochs and rhinoceros (Panagopoulou, et al. 2002-2004:340). No
carnivore remains were noted, and small game was not mentioned in the assemblage. Based
on a few fallow deer teeth, both adult and juvenile animals are present at the site
(Panagopoulou, et al. 2002-2004:341).
One of the most notable features of the Lakonis I project is the recovery of a
Neandertal lower left third molar from layer identified as Initial Upper Paleolithic (Unit Ia)
(Harvati, et al. 2003; Panagopoulou, et al. 2002-2004). This would be one of only a few sites
in Europe where human remains were found with a transitional stone tool industry, and in
this case it suggests that Neandertals were the makers of the IUP industry at Lakonis I
(Harvati, et al. 2009). Strontium levels in the tooth were analyzed using laser-ablation
PIMMS (Richards, et al. 2008). This analysis indicates that the enamel in the tooth formed
when the individual lived in an area with older, more radiogenic volcanic bedrock, as
77
opposed to the limestone cave in which it was found. The authors conclude that this
Neandertal lived as a child in an area at least 20 km away from Lakonis I (Richards, et al.
2008, but see Nowell and Horstwood 2009, who suggest that a different method of correcting
for isobaric inferences for Sr may actually indicate that the tooth formed locally).
Apidima
Apidima is a locality on the Mani Peninsula that includes four caves (Α, Β, Γ, Γ) that
contain Pleistocene deposits. Lithic artifacts and faunal remains were recovered, but the site
is better known for its human remains (Pitsios 1985, 1995, 1999). Two Middle Pleistocene
skulls were recovered Cave A, and an Upper Paleolithic burial of a 17-23 year-old female
was found in Cave Γ (gamma) (Ligoni and Papagrigorakis 1995). The skeleton in Cave Γ
was found laying on one side in the fetal position, with her head on a slab of stone, and the
burial was associated with a disk-shaped stone, animal bones, and perforated shell ornaments
(Cyclope neritea) (Karali-Giannakopoulou 1995:6-7; Pitsios 1999). The burial is thought to
date to about 30,000 BP based on associated lithics and the stratigraphic layer in which it was
found (Darlas 1995; Pitsios 1999). No reliable radiometric dates are available (but see
Lyritzis and Maniatis 1989, 1995).
A preliminary paper on the small lithic assemblage from Cave Γ suggests an
Aurignacian affiliation for the stone tools, though some Mousterian characteristics are also
apparent (Darlas 1995). Tsoukala (1999) presents a paleontological discussion of the faunal
remains at Apidima. In Cave Γ, she notes a wide range of small and large carnivores,
including leopard, lynx, wild cat, badger, and stone marten, as well as ibex, red and fallow
deer, Megaloceros, and Elephantidae. Based on the presence of juvenile carnivores, the
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author postulates that the cave was occasionally used as a den. Small animals, such as
rodents, birds, lagomorphs, and turtle bones are also present (Tsoukala 1999). Damage from
burning and butchery marks are present. Unfortunately, no quantitative values indicating
species frequency or taphonomic factors are available. In an analysis of the fauna of the four
caves, Lax (1995:133) determines that the faunal accumulations of Cave Γ (NISP = 7,383)
are largely natural, despite occasional burning (7.3%) and cut or chop marks on at least two
specimens.
Kephalari Cave
Kephalari Cave was exceptionally well excavated (including wet sieving of the
sediments) but never fully published. It lies in a network of caves above the small town of
Kephalari, about 6 km southwest of Argos (Felsch 1973; Reisch 1976). It was excavated
from 1972 to 1975 by the German Archaeological institute at Athens (Felsch 1973; Reisch
1976). The site contains a late Middle Paleolithic component, Upper Paleolithic layers, and
disturbed Neolithic deposits. Unfortunately, no radiocarbon dates are available for the site,
but diagnostic lithics indicate that it contains some kind of early Upper Paleolithic,
Aurignacian, Gravettian and Epigravettian components (Hahn 1984; Reisch 1976).
Occupations associated with the late glacial and the Last Glacial Maximum (LGM),
particularly the Gravettian, tend to be absent in other Peloponnesian sites (Adam 2007;
Karkanas 2010; Koumouzelis, Ginter, et al. 2001). If Kephalari Cave turns out to contain
LGM deposits, it has a lot of potential for adding to our understanding of the Paleolithic in
southern Greece. The lithics are currently under analysis by Dr. Gilbert Marshall. Worked
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bone is fairly abundant in the Kephalari Cave assemblage, and shell ornaments are common,
though no quantitative values are currently available.
A small portion of the Upper Paleolithic faunal remains were published in a brief
account by Reisch (1976). He notes a range of small animals in the assemblages, including
hedgehogs, bats, squirrel, and voles. Other small game species that were potentially of
economic importance at other Paleolithic sites in southern Greece include hare, tortoise,
partridge and fish (Reisch 1976). Small carnivores occur in the assemblages, including fox,
lynx, and wild cat, as well as some large hyena or leopard-sized carnivores. Ungulate species
identified include wild ass, wild pig, red deer, wild cattle, and wild goat (Reisch 1976). The
Kephalari Cave fauna is currently under study by the author and preliminary analysis
confirms the species list proposed by Reisch (1976). This new analysis also indicates that
large ungulates were much more common in the older layers, and hares, fish, and especially
partridge are found in large numbers in the later Paleolithic layers.
Franchthi Cave
Franchthi Cave, on the coast in the southeastern Argolid, is probably the most wellknown prehistoric site in Greece. It was excavated as part of a long-term interdisciplinary
study and resulted in an extensive publication series (e.g., Farrand 2000; Hansen 1991;
Jacobsen and Farrand 1987; Perlès 1987, 1991; Shackleton 1990; Stroulia 2010; Van Andel
and Sutton 1988; Wilkinson and Duhon 1991, in addition to many articles and book
contributions). The site preserves Upper Paleolithic through Neolithic deposits, with a
possible Middle Paleolithic component below the modern water table that could not be
excavated (Farrand 2000). The Campanian ignimbrite tephra, a well-known volcanic event
80
that occurred in Italy about 39,280 +/- 110 BP (40Ar/39Ar) (De Vivo, et al. 2001) is recorded
at the base of the Upper Paleolithic sequence; an Aurignacian occupation post-dates this
event, possibly by a few thousand years. Between about 33,000 BP and 25,000 BP, and after
15,000 BP, there was heavy use of the cave, punctuated by a few hiatuses (Farrand 2000).
The site is well known for its Neolithic component, but the Paleolithic and Mesolithic are the
focus of the review here.
Perlès (1987; 1991; 1999) divides the Upper Paleolithic and Mesolithic into nine
lithic phases based on stone tool abundance and technologies. Lithic phase I contains a small
sample but is typologically similar to the Aurignacian (Perlès 1987, 1999). A hiatus separates
lithic phases I and II; radiocarbon dates place the latter at 22,400 +/- 750 or 21,950 +/- 3,000
BP (Jacobsen and Farrand 1987). Lithic phase II mostly contains single and double backed
bladelets, though endscrapers and notches are also present (Perlès 1987, 1999). Lithic phase
III is stylistically similar to the preceding phases, with a higher proportion of single-backed
bladelets (Perlès 1987, 1999). A major hiatus spanning the LGM divides lithic phases III and
IV. Lithic phase IV dates to 14,850 +/- 800 BP (Jacobsen and Farrand 1987) and contains
backed bladelets, though there is no apparent reduction sequence for the manufacture of the
bladelets. The microburin technique appears at this time, as well as “La Mouillah points” in
the assemblage (Perlès 1987, 1999). In general, there is an increase in typological diversity in
lithic phase IV.
During lithic phase V, which dates to 13,200 +/- 300 BP (Jacobsen and Farrand
1987), the microburin technique was not used, and the first geometrics appear (Perlès 1987,
1999). Double-backed bladelets are common, and there is further diversification of tools,
including notches, denticulates and endscrapers. Perlès (1987) interprets this to be a different
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lithic tradition than those below. Four radiocarbon dates are available for lithic phase VI,
ranging between 12,600 +/- 1,450 and 12,100 +/- 750 BP (Jacobsen and Farrand 1987).
Production techniques in this phase are the same as those in lithic phase V, but there are
some typological changes. New geometrics appear in phase VI, and there is a further increase
in diversity of tool types. Also at this time obsidian from the island of Melos appears,
indicating seafaring in the Aegean (Perlès 1987).
There is a small hiatus between lithic phases VI and VII, which marks the beginning
of the Lower Mesolithic at Franchthi Cave. Seven radiocarbon dates were taken from the
Lower Mesolithic layers and span 10,725 +/- 475 and 10,275 +/- 375 BP (Jacobsen and
Farrand 1987). Stone tools in the Mesolithic differ drastically from those in the late Upper
Paleolithic in type and decline in quality in the later periods (Perlès 1991). Backed bladelets
and geometrics decrease in frequency in lithic phase VII. Most tools in the Lower Mesolithic
are laterally retouched and include notches, denticulates and endscrapers (Perlès 1991, 1999).
Lithic phase VIII represents the Middle Mesolithic and has five radiocarbon dates that range
between 10,000 +/- 400 and 9,550 +/- 300 BP, in addition to one questionable early date
(Jacobsen and Farrand 1987). There is a general paucity of lithic remains but an increase in
microliths at this time, which the author relates to tuna fishing (Perlès 1991). The Final
Mesolithic, lithic phase IX, dates to 9,215 +/- 215 BP (Jacobsen and Farrand 1987). The
lithic assemblage is quite similar to that during the Lower Mesolithic, with crude tools such
as notches and denticulates, with some arrowheads (Perlès 1991, 1999).
Bone tools are rare in the Franchthi assemblage. They are found in the late Paleolithic
layers (lithic phase VI) and in the Lower Mesolithic (lithic phase VII), though no extensive
publications are available on osseous tools at the site. The ornament assemblage from
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Franchthi Cave is large, with about 10,000 shell ornaments from the Upper Paleolithic and
Mesolithic deposits (Perlès and Vanhaeren 2010). The most common shells in the
assemblage are from Cyclope neritea, though Columbella rustica and Dentalium sp. are also
found (Perlès and Vanhaeren 2010). Interestingly, the shell ornaments at Franchthi Cave
were likely heat treated, possibly for aesthetic reasons. Perles and Vanhaeren (2010) note that
Cyclope neritea shells (both perforated and unperforated) display higher incidences of
burning, by several orders of magnitude, than other organic remains in the assemblage,
including animal bones and land snail shells, possibly for aesthetic purposes.
Faunal remains at Franchthi were initially studied by S. Payne (1975; 1982), who
determined that changes in the relative proportions of large mammals from about 25,000 BP
onward were related primarily to environmental but also economic change. He documented
an increase in small animals, such as birds, mollusks, and especially fish by the Mesolithic
(Payne 1975:128). The fish component in the Franchthi sequence shifted fairly dramatically
from the exploitation of brackish-water species such as eel, sea bream and gray mullet in the
Late Upper Paleolithic and Lower Mesolithic (ca. 13,020-10,230 BP) to open-water taxa
such as large bluefin tuna, grouper and barracuda in the Late Mesolithic (10,230-8,890 BP)
(Rose 1995). In this latter phase, the quantity of fish also increases in the faunal assemblage.
Marine mollusk exploitation shifted during this period, from shallow-dwelling Patella,
Monodonta and Gibbula at the end of the Upper Paleolithic to brackish-water Cyclope nerita
(for ornament-making) during the Lower Mesolithic and to Cerithium, a slightly deeper
water-dwelling gastropod, in the Late Mesolithic (Shackleton 1990).
Subsequent ongoing analyses of a large sample of the Franchthi faunas (NISP >
17,000) are presented by Stiner and Munro (2011). They identify eight major faunal zones
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that relate in part to changes in the local environment and resource intensification that may
have accompanied human population growth. These faunal zones do not altogether
correspond to lithic phases described above, so cultural designations are indicated. Upper
Paleolithic layers include Zone VIII (Aurignacian), which contains a diverse set of ungulates
dominated by red deer, and Zone VII (some Aurignacian/Gravettoid), which displays an
increase in wild ass and indicates a more open environment. The Epipaleolithic includes
Zone VI (Epigravettian) corresponds to an increase in small game exploitation, including
land snails, shellfish and coastal fishing. Zone V (Epigravettian) is marked by increased
cultural inputs, a further decline in ungulate diversity, and shift from tortoises to pond turtles.
Zone IV (Final Paleolithic) is dominated by red deer, and a notable disappearance of opendwelling taxa such as ground birds, aurochs and wild ass. The Mesolithic includes three
faunal zones: Zone III (Lower Mesolithic) has an increase in burned bone and a
corresponding increase in seed exploitation (Hansen 1991), Zone II (Mesolithic) contains
barracuda and tuna from open/deep water, and Zone I (Late Mesolithic) is dominated by
open or deepwater fished tuna (Stiner and Munro 2011). For the most part, shifts in large
terrestrial fauna (ungulates) correlate with changes in environmental conditions . Two,
consecutive trends of increasing dietary breadth are indicated by the small game data, the first within
an exclusively terrestrial context, and the second as marine habitats came into use through the end of
the Mesolithic. The intensity of the human occupations at this site increased in tandem with
intensified use of animal and plants (Stiner and Munro 2011).
Extensive botanical remains were recovered from Franchthi Cave, allowing for local
environmental reconstructions (see Chapter 3) as well as an evaluation of the contribution of
vegetation to the diet of foragers at the site (Hansen 1991). In the late Upper Paleolithic, from
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about 15,350 to 13,020 BP, edible species found in the site include pistachios, almonds,
pears, legumes (bitter vetch, lentils, vetch/vetchling), wild oats and barley (Hansen
1991:160). The late Upper Paleolithic/Lower Mesolithic transitional zone (cf. 10,495-10,230
BP) contains mostly pistachio, almond, pair and oats, with an increase in wild oats and barley
(Hansen 1991:161). The first part of the Late Mesolithic (10,230-9,645 BP) yielded few
botanical remains, but the period that follows until about 8,705 BP is similar in botanical
content to the late Upper Paleolithic/Mesolithic zone, with the addition of coriander (Hansen
1991:161). These data suggest the use of a wide suite of plants, thought it is difficult to
ascertain the quantitative role that each played in the diet.
Multiple burials and cremations were recovered from the Lower Mesolithic deposits.
Few Mesolithic human remains have been found in Greece, and Franchthi Cave is invaluable
in understanding mortuary practices from this period in the region. Seven fragments of
human bone were recovered in the Paleolithic layers, but the provenience for these remains is
not mentioned (Cullen 1995:274). At least eight burials were recovered in the Lower
Mesolithic layers: an infant, two young adults (male and female), three females and two
males, ranging in age from 20 to 45 years (Cullen 1995:275). At least two of the Lower
Mesolithic burials were also cremated. Only one possible intact burial was found in the Late
Mesolithic, a small collection of infant remains (Cullen 1995:272). Additionally, dozens of
isolated human bones and teeth were found throughout the Mesolithic layers, representing 725 individuals (nineteen is likely based on spatial distribution across the site) (Cullen
1995:278). The author suggests that human bone scattered across the site is the result of postdepositional disturbances, not as part of a funerary process (Cullen 1995:280).
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THE LATE MIDDLE PALEOLITHIC THROUGH MESOLITHIC
ARCHAEOLOGICAL SEQUENCE IN GREECE
One of the most striking features of the Middle Paleolithic through Mesolithic record
in Greece is the rarity of sites with a long or continuous occupation sequence that preserves
both lithic and faunal materials. Klithi, for example, has a large faunal assemblage, but the
site seems to represent a fairly short occupation span. Theopetra Cave, on the other hand,
contains a long stratigraphic sequence, but it has many long hiatuses and an overall lack of
faunal materials. Franchthi Cave, despite its many depositional hiatuses before the late Upper
Paleolithic, preserves by far one of the longest sequences of occupation with abundant lithic,
faunal and botanical remains. Of the sites discussed above, only Theopetra Cave and
Sarakenos Cave contain Middle Paleolithic, Upper Paleolithic and Mesolithic deposits; the
former is riddled with gaps and mixed deposits, and the extent of the Middle Paleolithic
layers of the latter are limited. Therefore, we are left to patch together the Late Pleistocene
and Early Holocene landscape in Greece from barely over a dozen archaeological sites.
Mousterian deposits are found at several Greek sites, including Asprochaliko,
Theopetra, Lakonis I, and Kalamakia. At all of these sites there appear to be technological
differences between the earlier and later Middle Paleolithic layers, though the patterns of
variation are not similar among the sites (e.g., the “micro-Mousterian” at Asprochaliko is not
found elsewhere). Middle to Upper Paleolithic transitional industries are absent in Greece,
with the possible exception of Lakonis, where a potentially stratigraphically sound layer
contains a mosaic of Middle and Upper Paleolithic production methods and tool types. It was
once thought that Theopetra Cave contained a transitional industry, but TL dates indicate that
the subject layer is, in fact, mixed. Aurignacian deposits are found at many Greek sites
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(Franchthi, Apidima, and possibly Kephalari), but only those at Franchthi are published.
Deposits dating to the same interval as the Eastern European Gravettian occur in Greece, but
the lithic industries are often called “Gravettoid” or “Gravettian-like”; cultural manifestations
of this time period seem to take on a regional variation (e.g., Asprochaliko, Kephalari,
Kastritsa and Grava) (see also Adam 2007).
The LGM is largely absent in the Greek archaeological record. It may be represented
at Kephalari Cave, but this is not certain as the deposits are undated. Some radiocarbon
ranges span this period, such as at Kastritsa, but at most sites the LGM corresponds to a
hiatus (Franchthi, Theopetra). It is unclear if Greece was depopulated during the LGM, if the
sites did not preserve, or if people were concentrated in coastal areas that are now
underwater. Late Upper Paleolithic sites are comparatively common, and often contain
Epigravettian components (e.g., Theopetra, Klithi, Boïla, Sarakenos Cave, Kephalari and
Franchthi). Mesolithic deposits are found at Theopetra, Boïla, Sarakenos Cave, the Cave of
Cyclope and Franchthi Cave; the latter two contain extensive archaeological materials.
Another feature of the Greek Paleolithic is that lithic raw material sources appear to
be primarily local and often of low quality. This may seem to be the case because no
comprehensive provenance studies have been conducted, and because many raw materials
used in the Paleolithic are not conducive to sourcing. The situation changes at the end of the
Upper Paleolithic and in the early Mesolithic when non-local materials, particularly obsidian
from Melos, are found at sites in Peloponnese (Franchthi Cave), as well as in the northern
Aegean (Cave of Cyclope). Organic artifacts, including bone points, fishhooks and
ornaments, appear during the Upper Paleolithic in Greece and proliferate in the Mesolithic at
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some sites. A few questionable early pieces were recovered, as well as the retouched shell
scrapers at Kalamakia, but these are fairly rare in the Middle Paleolithic.
Faunal remains are difficult to compare among the sites discussed above, mostly
because of preservation issues or an altogether lack of published quantitative data. The large
game component of Paleolithic Greek sites may be largely reflective of local environments
rather than deliberate variation in prey choice by human hunters. The dominance of ibex and
chamois in the faunas of Klithi, for example, correlate with the mountainous region in which
the site is located. At sites such as Apidima, Asprochaliko, Kalamakia, Lakonis, megafauna
that later go extinct (Pleistocene rhinoceros and elephants) occur in the lower layers.
Carnivores are common in many but not all of the faunal assemblages discussed above (e.g.,
Kalamakia, Theopetra, Apidima). Without rigorous taphonomic studies of these assemblages,
it is unclear the extent to which carnivores shaped at least some of these archaeofaunas (see
Chapter 4). Diachronic change in faunal assemblages is also difficult to ascertain from the
sites discussed above with the exception of Franchthi Cave, because of small samples and the
limited nature of temporal sequences.
One robust trend comes from Greek coastal sites dating to the Late Pleistocene and
Early Holocene (e.g., Franchthi Cave, Cave of Cyclope and Sidari), where there is an
increase in use of marine resource. Contemporary terrestrial Mesolithic assemblages are
essentially absent in the Greek record (the faunal sample from this time period is
exceptionally small at Theopetra cave). It is unclear at this point if this represents a general
movement of human populations to coastal areas, or if it is a bias in the archaeological record
in both the location of excavated Mesolithic sites and the preservation of terrestrial faunas at
inland sites that do not contain Mesolithic layers.
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While terrestrial mobility may have declined towards the end of the Pleistocene, there
are indication of increased mobility on marine waters via innovations in seafaring,
particularly during the Mesolithic period. This is evidenced by tuna fishing, and the
appearance Melos obsidian on the mainland and on other Aegean islands. The inclusion of
fish and marine mollusks at coastal sites indicates a widening of the diet breadth of Early
Holocene foragers in this region.
Klissoura Cave 1 has the potential to fill many of the gaps in the regional record
discussed above. The site is 12 km inland, and contains Middle Paleolithic through
Mesolithic deposits. It also preserves an Early Upper Paleolithic layer and thick Aurignacian
deposits unlike any found in Greece. There are some depositional hiatuses, including during
the LGM, but there are ample, well-preserved lithic, faunal and botanical remains in almost
all of the cultural layers represented at the site. I now turn to a discussion of Klissoura Gorge
and its caves, specifically Klissoura Cave 1.
KLISSOURA GORGE AND KLISSOURA CAVE 1
Klissoura Gorge is part of a valley that stretches for approximately 3 km, connecting
the Berbati Valley and Argive Plain in northeastern Peloponnese (Figures 2.1, 2.3). The
gorge narrows to 500 m wide in the area that contains several karstic cave sites of the same
name. The valley formed in Triassic limestone and is drained by the Berbatiotis River
(Koumouzelis, Ginter, et al. 2001). Thirty-five caves and rock shelters were systematically
surveyed in the gorge, six of which contain surface archaeological materials, while a handful
of others preserve stratified sequences (Koumouzelis, et al. 2004). The gorge was initially
surveyed and recorded during the Berbati-Limnes Archaeological survey in 1988-1990
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Figure 2.3 Photograph of Klissoura Gorge, facing southwest.
(Runnels 1995a). A joint project between the Ephory for Paleoanthropology and Speleology
in Athens and the Institute of Archaeology at the Jagiellonian University in Kraków tested
Klissoura Caves 4 and 7 in 1993, while systematic excavations began at Klissoura Cave 1 in
1994 (Koumouzelis, Ginter, et al. 2001; Koumouzelis, Kozlowski, et al. 2001; Koumouzelis,
et al. 1996). Today, the gorge lies roughly 12 km inland; in the past the closest marine
shorelines were slightly further away (Van Andel and Shackleton 1982) (Figure 2.4).
Three 1x1 meter test trenches were excavated in Klissoura Cave 7, yielding three
stratigraphic layers with a few hundred lithic artifacts in each layer (Koumouzelis, et al.
2004). A radiocarbon date on calcium carbonate yielded a date of 14,850 +/- 700 BP, but the
authors contend that the true age is slightly younger (Koumouzelis, et al. 2004:35). Surface
finds included some tools and flakes attributable to the Mesolithic and Upper Paleolithic, and
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Figure 2.4 Comparison of modern landmass (grey fill) to LGM shorelines (black line and white fill,
indicating land inundated today) and paleolakes (black fill). Adapted from Petit-Maire et al. (2005).
the authors propose that stone tools found in the excavated layers correlate stylistically with
lithic phases V and VI at Franchthi Cave, which are late Upper Paleolithic (Perlès 1987).
At Klissoura Cave 4, two 1x1 meter test trenches were excavated, one of which
contained primarily mixed sediments. The second trench includes several layers of mixed
Neolithic and Paleolithic artifacts, overlying several intact late Upper Paleolithic deposits
(Koumouzelis, et al. 2004). The authors postulate that these layers also correspond to the late
UP at Franchthi Cave, specifically lithic phases V and IV (Perlès 1987). A bone sample from
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one of the lower layers of Klissoura Cave 4 dates to 13,600 +/- 750 BP (Koumouzelis, et al.
2004:43).
The initial excavations at Klissoura Cave 1 (Figure 2.5) established the presence of
Holocene layers containing Mesolithic flake industries and Late Pleistocene layers that
include Aurignacian-like industries. Subsequent excavations revealed a deep series of
deposits that include thick Upper and Middle Paleolithic components (Koumouzelis, Ginter,
et al. 2001; Koumouzelis, Kozlowski, et al. 2001; Sitlivy, et al. 2007). The site was
excavated in 1x1 meter units (Figure 2.6) in 5 cm arbitrary levels, and all sediments were
screened. Excavations at the site began in 1994 and concluded in 2006.
Figure 2.5 Photograph of Klissoura Cave 1, facing northwest.
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Figure 2.6 Plan map of Klissoura Cave 1. Adapted from Karkanas (2010).
Stratigraphy and Dating of Klissoura Cave 1
In total, about a six meter deep trench was excavated from Klissoura Cave 1, with
twenty-one stratigraphic layers and various sub-groups in many of the layers. Table 2.1 lists
the excavated layers along with their cultural designations as defined by stone tool
typologies, radiocarbon date ranges, and depositional histories (Kaczanowska, et al. 2010;
Kuhn, et al. 2010; Sitlivy, et al. 2007). Certain stratigraphic units were grouped together to
improve sample sizes for the faunal study. Middle Paleolithic layers are grouped by
sedimentary units that formed in similar or related depositional environments (Sitlivy, et al.
2007; Karkanas personal communication 2009), while the Upper Paleolithic and later layers
correspond to changes in material culture (Kaczanowska, et al. 2010; Koumouzelis, Ginter, et
al. 2001; Koumouzelis, Kozlowski, et al. 2001). Koumouzelis, Ginter, et al. (2001) list a first
series of radiocarbon dates from the Upper Paleolithic and later layers of the excavation.
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However, these dates were taken mainly on ash-derived carbonate fraction in the sediments,
which likely underwent some recrystallization (Koumouzelis, Kozlowski, et al. 2001; Kuhn,
et al. 2010). Therefore, most of these initial dates probably represent minima and are not
presented here. A new series of AMS radiocarbon dates were recently established for
Klissoura Cave 1, based almost entirely on carbon, typically wood charcoal (Kuhn, et al.
2010). Pre-treatment techniques for cleaning samples obtained from the early Upper
Paleolithic and late Middle Paleolithic layers include the ABOX technique. The ABOX
"super-clean" pre-treatment method was developed by Bird et al. (1999) and employs wet
oxidation and step heating of samples that are subsequently processed in a special vacuum
line (Pigati, et al. 2007). This method is well-equipped to deal with samples that date to
Table 2.1 Stratigraphic layers for Klissoura Cave 1 and corresponding cultural units based on stone
tool types. Layer groupings are used throughout the dissertation for analyzing faunal remains. From
Kaczanowksa et al. 2010, Karkanas 2010, personal communication 2009, Kuhn et al. 2010.
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beyond the range of traditional radiocarbon dating, and samples that contain a small amount
of carbon.
Kuhn et al. (2010) present the full list of dates taken at Klissoura Cave 1; an abridged
version of this is provided in Table 2.2. Certain layers are excluded (specifically 6, 6a, 7, and
VI) because analysis of the artifacts shows these layers to be mixed. I also exclude dates
from areas with ambiguous contacts between layers; for example, it is not clear if a date from
layer “IV/V” is lower Aurignacian or Early Upper Paleolithic. Four radiocarbon dates come
from the Middle Paleolithic layers, ranging from roughly 53,600 to 62,300 BP (Table 2.2,
note the large error ranges). The three oldest dates are considered minimum ages for the two
Middle Paleolithic layers from which they were derived.
Radiocarbon dates from the Early Upper Paleolithic (Uluzzian) layer V remain
problematic, which is unfortunate because this layer represents a rare industry in Greece.
Most of the dates are either anomalously young, infinite, or of questionable provenience
(Table 2.2). Kuhn et al. (2010:41) propose two scenarios for the age of the layer. The first is
that the older date (43,841 +/-764 BP) is correct, as are two similar dates from the underlying
mixed layer VI, which would indicate a hiatus of 6,000-7,000 years separating the EUP and
lower Aurignacian layers. The second possibility is that the ~40,000 BP dates are intrusive
from the late Middle Paleolithic, and the EUP layer V was deposited sometime between
33,000 and 40,000 BP. Recent analysis of a micro-tephra from the interface between layer IV
and V indicates that it is the Campanian Ignimbrite (Karkanas, personal communication
2011), which is also found at Franchthi Cave and dates to 39,280 +/- 110 BP using 40Ar/39Ar
(De Vivo, et al. 2001). This is consistent with the first scenario proposed by Kuhn et al.
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Table 2.2 Radiocarbon dates from Klissoura Cave 1. From Kuhn et al. (2010).
96
(2010), where there is a large hiatus between the EUP and Aurignacian. It seems likely,
therefore, that the Uluzzian of layer V is greater than 39,000 BP.
Some anomalously young dates for the rest of the Upper Paleolithic layers are noted
in Table 2.2, but for the most part the dates from these layers appear sound. The Aurignacian
at Klissoura Cave 1 was probably deposited between 35,000 and 37,500 BP (Kuhn, et al.
2010:40). The Mediterranean-backed bladelet industry (III’), which is the uppermost of two
distinctly non-Aurignacian layers (see below), yielded a date of 35,381 +/- 690 BP (Table
2.2), as well as an anomalously young date. No dates in the new series address the ages of the
Epigravettian (IIa-d) and Mesolithic (3-5a) layers, but some are presented in Koumouzelis,
Kozlowski et al. (2001). These are radiocarbon dates on sediment carbon fraction that
yielded ages of 17,150 +/- 600 and 10,350 +/- 800, respectively.
Stratigraphy
The depositional history of Klissoura Cave 1 is fairly well understood.
Geoarchaeological analyses and micromorphology of the Middle Paleolithic layers are still
being studied (but see Sitlivy, et al. 2007 for preliminary results), but the analysis of the
Upper Paleolithic layers are completed (Karkanas 2010; Karkanas, et al. 2004). The Middle
Paleolithic sediments contain a much higher geogenic input, as opposed to the heavily
anthropogenic signature of the Upper Paleolithic layers (Sitlivy, et al. 2007).
The lowest sedimentary layer at Klissoura Cave 1 is XXI, which is only found toward
the eastern wall of the shelter (Sitlivy, et al. 2007). There is a large disconformity between
this layer and the overlying Middle Paleolithic deposits, suggesting a major environmental
event at this time. The faunal component from XXI is also different from overlying layers
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and is composed primarily of microfauna and small bird remains. Because the fauna was
determined during analysis to be non-anthropogenic, it is not included in this study. The
overlying layers include XX through XVI (Table 2.1). Sediments in these layers are finegrained, with a high degree of post-depositional chemical alteration, especially in layer XX
(Sitlivy, et al. 2007:4). The lower deposits slope toward the entrance of the site, and the
authors note that layer XVIII is especially rich in lithic and bone artifacts. Sitlivy et al.
(2007) describe a difference between the stratigraphically higher (VII-XIV) and lower (XVXX) Middle Paleolithic sediments: the higher layers are composed of stone-rich, reddish
clay, and the lower layers contain fine-grained grey silty and ashy sediments. The authors
believe that the difference between the upper and lower Middle Paleolithic layers is
explained by a collapse of the back chamber of the cave during the formation of layer XV.
This collapse created a chimney that allowed rainwater, terra rossa sediments, and roof fall
to wash into the cave (Sitlivy, et al. 2007:4).
The stratigraphically higher Middle Paleolithic layers (VII-XV) are composed of
brownish-red fine-grained sediments with stony deposits at the back of the shelter, indicating
a higher input of clay and clastic materials (Sitlivy, et al. 2007:3). Layers XI-XIV contain
alternating clay-rich layers and thin, anthropogenic burned layers. There is an erosional
depression at the front of the shelter that truncates layers XI and XII (Sitlivy, et al. 2007:3).
Layer IX contains very few faunal remains, but this layer along with layer X contain burned
remains cemented by surface water. An erosional event formed a trough in layer X that was
subsequently filled by either natural sediments or anthropogenic materials (Sitlivy, et al.
2007:3-4). Layers VI through VIII contain clay-rich sediments and an increasing proportion
of anthropogenic inputs, including burned lenses. It was determined by the analysis of the
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lithic and ornament assemblages that layers VI and VII are mixed in some parts of the site,
containing both Middle and Upper Paleolithic materials (Sitlivy, et al. 2007; Stiner 2010;
Stiner, et al. 2010); consequently, layers VI and VII were excluded from the faunal analysis.
Based on geoarchaeological and micromorphological analyses, Karkanas (2010)
divides the Upper Paleolithic and later cultural layers in Klissoura Cave 1 into depositional
facies that are grouped into a stratigraphic sequence. An in-depth description of the facies is
presented in Karkanas (2010). In general, the Upper Paleolithic deposits are defined by an
extremely high anthropogenic input. Anthropogenic inputs decrease somewhat in the
Epigravettian (IIa-d) and Mesolithic (3-5a) layers, which are dominated by clastic sediments
(Karkanas 2010:32).
The Early Upper Paleolithic or Uluzzian layer V is significant because it is one of few
EUP deposits found in Greece. It is composed of dark gray clayey silt with reworked and in
situ burned features in some areas. Although layer V is diffuse in some parts of the
excavation, it is intact in the middle and southern parts of the site (Karkanas 2010:30). The
deposits in layer V represent a similar sedimentary environment as those that formed layer
IV, though based on recent dating evidence discussed above, there is reason to suspect that a
hiatus of several thousand years separates the formation of the two layers (Kuhn, et al. 2010).
Layer IV, the lower Aurignacian, contains many burned features and deposits,
including numerous clay-lined hearths (see below) (Karkanas 2010; Karkanas, et al. 2004;
Pawlikowski, et al. 2000). The sediments are gravelly silts and are light in color due to a high
wood ash content. Clay-lined hearth structures were not found in the southern units of the
excavation, near the entrance of the shelter; rather the sediments in this area are dominated
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by homogenous reddish-brown clay (Karkanas 2010:30). A linear or clustered arrangement
of large stones representing a possible structure are found in this area.
The layer III series is filled with burned deposits from reworked hearths, intermingled
with undisturbed deposits. This layer also contains clay-lined hearth structures, along with
some gravel-rich clusters (Karkanas 2010:30). Many of the stratigraphic layers included in
this sequence are horizontally discontinuous, and were not necessarily deposited
simultaneously. Karkanas (2010) does note, however, that units within the layer III series can
be grouped into a coherent stratigraphic order: IIIe-g are the oldest and are overlain in
succession by IIIb-d, III", III', and finally III. This is relevant to the discussion of stone tool
industries discussed below, as the bottom-most two layer groups (IIIe-g and IIIb-d) are
Aurignacian deposits, and III’ and III” are assigned to distinct non-Aurignacian
Mediterranean blade or backed bladelet industries (Kaczanowska, et al. 2010; Koumouzelis,
Ginter, et al. 2001; Koumouzelis, Kozlowski, et al. 2001). The contact between layer III’ and
the Epigravettian layers is locally diffuse. Some freeze-thaw action is apparent in certain
parts of layer III’ but not in any other layer of Klissoura Cave 1 (Karkanas 2010:34).
An anthropogenic ditch or intrusion of mixed deposits (6-7a) cuts into layers III’ and
III” in certain parts of the site. This layer is often referred to in earlier publications (e.g.,
Karkanas, et al. 2004; Koumouzelis, Ginter, et al. 2001; Koumouzelis, Kozlowski, et al.
2001; Starkovich 2009) as part of the Upper Paleolithic sequence, but it is now known to be
mixed based on the lithic results and is excluded from this study.
The Epigravettian layers (IIa-d) are composed mostly of brown, clay-rich deposits,
though in certain areas there are reworked burned remains and sorted, laminated sediments
(Karkanas 2010:28). Clusters and lines of limestone rocks representing deflated surfaces (i.e.
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hiatuses) are common. Some bioturbation from insects was noted in the Epigravettian layers
(Karkanas 2010:34).
The youngest layers include the modern surface and disturbed Bronze Age deposits
(1-2), through the partly intact Mesolithic deposits (3-5a). The uppermost parts of these
layers have numerous recent pits and evidence of animal trampling. The Mesolithic layers (35a) are composed of clay-rich deposits with lenses of loose ash and shell and laminated
geogenic layers (Karkanas 2010:28).
Lithic Assemblages
Tens of thousands of stone tools or lithic fragments were recovered from the Middle
Paleolithic at Klissoura Cave 1, and over 130,000 from the Upper Paleolithic and later layers
(Kaczanowska, et al. 2010; Sitlivy, et al. 2007). Most of the assemblages are composed of
chips or debitage, followed by flakes and tools (Kaczanowska, et al. 2010; Koumouzelis,
Ginter, et al. 2001; Sitlivy, et al. 2007). The site does not preserve an uninterrupted sequence
during the span of occupation (Kaczanowska, et al. 2010); rather there are several hiatuses in
the sequence, as discussed previously.
The Middle Paleolithic lithic assemblages from Klissoura Cave 1 are still under
analysis, but preliminary results from a large sample (n = 37,922) are presented in Sitlivy et
al. (2007, see also Koumouzelis, Kozlowski et al. 2001). Proportions of raw material types
are similar throughout the sequence, with a dominance of radiolarite followed by flint. In
terms of rare raw material types, limestone and quartz are more common in the lowermost
and middle part of the MP sequence, and chalcedony occurs in the uppermost layers (Sitlivy,
et al. 2007:6).
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The authors find many other similarities in the Middle Paleolithic lithic assemblages
of Klissoura Cave 1, and a high degree of homogeneity throughout. In general, Middle
Paleolithic tools and debitage fragments are quite small in all layers, though blanks are
slightly larger in the lowest units (Sitlivy, et al. 2007:10). The earliest phases of raw material
testing and removal of the cortex are absent at the site, suggesting that the occupants brought
pre-tested raw materials to the cave for tool production (Sitlivy, et al. 2007:9). A variety of
reduction strategies are found in all layers, including unidirectional or discoidal and
centripetal methods, and the frequency of Levallois technologies is low throughout
(Koumouzelis, Kozlowski, et al. 2001; Sitlivy, et al. 2007). The tool assemblages are
primarily Mousterian, though some Upper Paleolithic elements are apparent in the younger
layers, indicating some mixing in certain parts of the site.
Some subtle differences are also apparent among the Klissoura Cave 1 Middle
Paleolithic lithic assemblages. The proportion of blades and bladelets is much higher in the
lowest layers (XVIII, XIX and XX) (Sitlivy, et al. 2007:8). The authors note that volumetric
production methods typically associated with Upper Paleolithic industries were used to make
blades in the lower layers, along with other reduction strategies, though Upper Paleolithic
tools are not present in the older Mousterian layers (Sitlivy, et al. 2007:12). By far the most
common tools in the Middle Paleolithic assemblages are side scrapers, though their
frequencies vary slightly with time. They occur in the smallest numbers in the latest (VII and
VIIa) and earliest (XVIII and XIX) Middle Paleolithic layers, and they peak in the middle of
the sequence (XIII) (Sitlivy, et al. 2007:11). Mousterian points are documented in all layers
except VIIa, and they are the most common in the lower layers (XVIII and XIX). Two
different styles of points are recorded, elongated points and short massive types. Most of the
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variability in point form is found in the lowest layers (XVIII, XIX and XX) (Sitlivy, et al.
2007:12). Upper Paleolithic tool types, including endscrapers, burins, perforators and
splintered pieces are found in the youngest Middle Paleolithic layers (VII, VIIa and VIII),
and completely disappear by layer X (Sitlivy, et al. 2007:12). Although the contents of layers
VII, and especially VIII, are essentially Middle Paleolithic, the possibility of low levels of
mixing from the early Upper Paleolithic cannot be excluded.
Sitlvy et al. (2007) conclude that Klissoura Cave 1 is different from other Greek
Middle Paleolithic sequences. Specifically, the authors note that there is no “microMousterian” overlying Levallois/Mousterian layers such as that found at Asprochaliko
(Bailey, et al. 1983). Also, though there are small numbers of bifacial tools in the lowest MP
layers, these are absent in the later layers at Klissoura Cave 1, suggesting a different
evolution of stone tool technologies in southern and northern Greece (Sitlivy, et al. 2007:14).
The Early Upper Paleolithic layer (V) in Klissoura Cave 1 is similar to the EUP
industries of Italy and has thus been defined as Uluzzian (Kaczanowska, et al. 2010;
Koumouzelis, Ginter, et al. 2001). This is significant as it is the first published Uluzzian
occupation in Greece and represents a rare and exceptionally old Early Upper Paleolithic
industry in the country. The sample is small compared to the later layers (n = 4,228). Flakes
are common in the assemblage, as are blades and tools. Radiolarite is the most common raw
material, followed by local flint (Kaczanowska, et al. 2010). Diagnostic artifact types in this
layer include arched backed blades and convex truncations along with splintered pieces;
endscrapers, retouched blades, and notched denticulated tools also occur but are scarce
(Kaczanowska, et al. 2010; Koumouzelis, Ginter, et al. 2001).
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A thick series of Aurignacian deposits was excavated at the site, with assemblages
defined by micro-retouched bladelets and carenate cores or endscrapers (Kaczanowska, et al.
2010; Koumouzelis, Ginter, et al. 2001). Based only on their stratigraphic position, they are
divided by Karkanas (2010) into the lower (IV), middle (IIIe-g) and upper (IIIb-d)
Aurignacian units (though Kaczanowska, et al. 2010 divide the latter two as IIId-g and IIIa-c;
this is not problematic in terms of the fauna as almost all analyzed material came from IIIc,
IIIe, or IIIg). In the Aurignacian series, as with the EUP, the most common raw material type
is radiolarite, followed by flint. In general, the composition of tools types in the series is
similar, with end scrapers being the most common, followed by denticulated-notched pieces,
retouched flakes, burins, side scrapers, and blades/retouched bladelets, among others
(Kaczanowska, et al. 2010). The bulk of the Klissoura Cave 1 lithics are found in the lower
Aurignacian layer IV (n = 63,837), a quarter of which were recovered from within a rocklined structure (see below) (Kaczanowska, et al. 2010). About half of the recorded Upper
Paleolithic hearths were also found in this layer (n = 55). Aurignacian layer IIId-g contains a
large sample as well (n = 28,625), whereas layer IIIa-c yielded the smallest sample of the
series (n = 5,623). Thirteen lumps of ochre were recovered in the middle Aurignacian layers,
and two from the upper (Kaczanowska, et al. 2010), though their sizes are not mentioned.
Above the Aurignacian layers there are two technically distinct but unnamed
Mediterranean Upper Paleolithic backed bladelet or blade industries. The older of the two
layers (III”) is referred to in this presentation as the non-Aurignacian Upper Paleolithic, and
the younger (III’) is called the Mediterranean backed bladelet industry. The non-Aurignacian
Upper Paleolithic is a relatively small lithic assemblage (n = 2,935). Flakes are much more
common than blades in this layer, and tools include endscrapers, denticulates and notched
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tools, retouched flakes and side-scrapers (Kaczanowska, et al. 2010). Layer III” is different
from the underlying Aurignacian because of the importance of flakes in blank production,
few cores for bladelet production, and few carenate cores or endscrapers.
The Mediterranean backed bladelet industry layer III’ is distinct because of the
importance of backed bladelets, microretouched bladelets, and truncations (Kaczanowska, et
al. 2010). The lithic sample from this layer is moderately sized (n = 5,943), and most of the
pieces are radiolarite followed by flint. The most common large tool type is again endscrapers, though backed bladelets and microliths occur in much higher frequencies in layer
III’ than in other layers at the site, followed by retouched flakes (Kaczanowska, et al. 2010).
Kaczanowska et al. (2010) suggest that this may represent one of the oldest backed bladelet
industries in the Mediterranean Basin, and they compare the tools in layer III’ to those found
in the Asprochaliko sequence.
After a stratigraphic hiatus representing the LGM, there is a comparatively limited
Epigravettian deposit (IIa-d, n = 6,312) containing an industry that is defined by backed
blades with ventral retouch, para-geometric forms and shouldered points (Kaczanowska, et
al. 2010; Koumouzelis, Ginter, et al. 2001). Most of the lithic materials are made of
radiolarite, followed by flint. Backed tools, microlithics and truncations are common,
followed by endscrapers, and blades with ventral retouch.
The most recent major stratigraphic layer at Klissoura Cave 1 is a Mesolithic (3-5a)
deposit with 6,812 lithics (Kaczanowska, et al. 2010). As with the earlier layers, radiolarite is
common followed by flint, though two obsidian artifacts are also found (Kaczanowska, et al.
2010; Koumouzelis, et al. 2003). Given that obsidian from the island of Melos appears
during the early Mesolithic at Franchthi Cave, it is possible, but not certain, that the obsidian
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from Klissoura Cave 1 is also from Melos. In the Mesolithic layers there is a much lower
proportion of chips and lithic debris as compared to tools, so it seems likely that many tools
found in the assemblage was manufactured off-site (Kaczanowska, et al. 2010). Tools
include endscrapers, side scrapers, retouched flakes and blades, microlithis, and geometric
forms, among types rarer in the assemblage. The authors note that the Klissoura Cave 1
Mesolithic is unlike the industries found at Franchthi Cave (Kaczanowska, et al. 2010;
Koumouzelis, et al. 2003).
Botanical Remains
Phytoliths and macrobotanical remains were studied in the Upper Paleolithic and later
layers of Klissoura Cave 1; analyses of the Middle Paleolithic layers are ongoing. The
contribution of these studies to local paleoenvironmental reconstructions are discussed in
Chapter 3. Phytolith samples were collected from the Aurignacian and later layers, from the
tops of clay-lined hearth structures (see below) and other hearths, and from the surrounding
matrix (Albert 2010). Importantly, few phytoliths were found in hearth samples in any of the
layers. Albert (2010) suggests that this finding indicates that the leaves and stems of
monocotyledonous taxa and wood or bark were the most common fuel sources, as wood and
bark contain far fewer phytoliths than grasses, particularly in certain Mediterranean species
(Albert and Weiner 2001). Phytoliths from grasses with edible seeds were found outside of
hearth areas in the Aurignacian series, as well as in the two layers that contain backed
bladelet industries (III’ and III”). Sedges, which are high in starch and protein content and
also useful materials for fiber-working, were found especially within the layer III series, but
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also in the lower Aurignacian (IV) and Epigravettian (IIa-d) (Albert 2010). No phytolith
samples are available for the Mesolithic (3-5a).
Water-flotation was undertaken on samples from hearths and sediments from selected
excavation units in order to collect wood charcoal (see Ntinou 2010 for a full description of
methodology). In general, large pieces wood charcoal are rare in hearths, occurring in only
twenty-nine of over a hundred of hearths, though microscopic charcoal is found throughout
the layers in all parts of the site (Karkanas 2010). Further, there were no higher incidences of
charcoal in clay-lined or stone-delineated hearth structures. Ntinou (2010) postulates that the
lack of macroscopic charcoal is due to the intense use of hearths, with complete combustion
of plant materials or cleaning events after each hearth was used. The most common plant taxa
used as fuel at the site include maple, deciduous oak, and Prunus sp., and deciduous oak. The
proportions of these fuel sources vary among hearths within and between layers (Ntinou
2010).
Features
Among the most unique features at Klissoura Cave 1 are the dozens of clay-lined
hearths in the Aurignacian and overlying Upper Paleolithic deposits (Karkanas, et al. 2004;
Kot 2009; Pawlikowski, et al. 2000). These features represent some of the earliest recorded
uses of modeled clay in the world (Pawlikowski, et al. 2000). Over a hundred well-preserved
hearths were encountered in the archaeological sequence of Klissoura Cave 1, in all layers.
However, over fifty clay-lined hearths are confined to the layers that contain the nonAurignacian backed bladelet industries (III’ and III”) and the underlying Aurignacian series
(IIIb-d, IIIe-g, IV) (Karkanas, et al. 2004; Pawlikowski, et al. 2000). The earliest clay-lined
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features at Klissoura Cave 1 date to between 35,000 and 37,500 BP, which predates the
Gravettian clay figurines from Dolni Vestonice (Vandiver, et al. 1989) by over five thousand
years. During excavation, these orange, basin-shaped features were easily recognized and are
obvious in profile (see images in Karkanas 2010; Karkanas, et al. 2004). Later clay-lined
hearths are often superimposed on earlier ones. Most of the clay-lined and unlined hearths
range between 30 and 40 cm in diameter, and a few of the clay-lined features have rims
(Karkanas, et al. 2004:515). Generally, the clay-lined features are covered by ashy lenses
filled with charcoal fragments, lumps of soil, and burned lithics and bone. Analyses of
micromorphology samples from the hearths and from an incidental mudflow in the cave
indicate that the hearths did not form in-situ via a natural process (Karkanas, et al. 2004:518).
Rather, the clay was brought to the site, probably from the floodplain in the base of the
gorge, and then wetted and pooled into the basin-like shape. Fourier transform infra-red
spectrometry (FTIR) and differential thermal analysis (DTA) performed on samples from the
clay-lined hearths indicates that they were heated between 400 and 600 °C (Karkanas, et al.
2004, but see also Pawlikowski et al. 2000, who suggest that at least one hearth was fired
above this temperature based on the degradation of the mineral kaolinite). It is not known if
the clay hearths were hardened with fire in anticipation of their first use or instead hardened
as the result of use.
Kot (2009) presents the results of experimental studies on the formation of the claylined hearth structures at Klissoura Cave 1 and proposes five possible uses for the features:
long-term ember storage, exposed cooking or roasting meat and plant foods, water boiling,
surfaces for holding food during burial with hot ashes or embers, or easily cleaned surfaces
for reuse as hearths (see also Meignen, et al. 2001). Drying or curing meat and skin are also
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possibilities for clay-lined hearth use (Ntinou 2010). Taking the time to create such structures
indicates longer term or more intensive site use, which is attested to by the influx of artifacts
and anthropogenic sediments into the site during these periods of occupation. Karkanas et al.
(2004) suggest that the small clay-lined structures were satellite hearths around a larger flat
(simple) hearth. They also note that starch and seed grass phytoliths were found in a sample
taken from the basin of one of the clay hearths, so it is possible that they were used on some
occasions for cooking or parching plant materials. In addition to the clay-lined structures,
five hearths are rock-lined and four are associated with stones (Ntinou 2010). Nearly all of
these structures are found in the lower Aurignacian (IV), though one is from the Early Upper
Paleolithic (V), and one other from the middle Aurignacian (IIIe-g).
Another interesting feature, a circular man-made rock alignment, was noted in the
lower Aurignacian (IV) (Koumouzelis, Ginter, et al. 2001; Stiner 2010; Stiner, et al. 2010).
This structure is apparent in plan maps (see Chapter 7) and is defined by a large circle of
limestone rocks, dark staining inside the circle, a lack of hearths, and a concentration of
ornaments and lithic debris within the structure. The structure takes up four excavation units,
giving it a maximum diameter of about 1.5-2 meters (Koumouzelis, Ginter, et al. 2001).
Ornaments
Shell ornaments are common in the Upper Paleolithic Klissoura 1 sequence, with
over 1,500 recovered (Stiner 2010). Ornaments are found in all UP and later layers, though
their frequencies change dramatically. The majority of the ornaments are from the lower
Aurignacian (IV), followed by the middle Aurignacian (IIIe-g). Ornament frequencies are
quite low in the Epigravettian and Mesolithic layers, partially due to small sample sizes for
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all cultural materials (Stiner 2010). The author suggests that declines in mollusk species
richness after the middle Aurignacian (IIIe-g) likely corresponds to the expansion of lagoon
formations in the western Argolid. Similar to other Paleolithic sites in Greece, Cyclope sp. is
the most common species in the assemblages overall, followed by Columbella rustica. Stiner
(2010) notes that the sample size for the EUP layer is small, but that Dentalium is
particularly common, not unlike that situation in some Italian Uluzzian sites (e.g., Palma di
Cesnola 1966).
Foragers occupying Klissoura Cave 1 collected mainly vacant shells from the marine
littoral, along with exploited fossil deposits and some freshwater sources. They
disproportionately selected the shells of predatory and scavenging carnivorous taxa for
ornaments, possibly because of their scarcity or aesthetic value, a phenomenon that is found
elsewhere in the Mediterranean (Stiner 2003a, 2010). Ornaments at the site display a narrow
size range: 1.3-1.4 cm in the Upper Paleolithic, and 1.4-1.7 cm in the later layers. Naturally
red Clanculus and striped Theodoxus shells were enhanced with red ochre (Stiner 2010).
Unlike the situation at Franchthi Cave, burning frequencies on ornaments at Klissoura Cave 1
are probably from random, accidental burning. Cord wear is documented on many of the
ornaments, indicating they arrived at the site already strung as beads or fastened to organic
materials. A final observation about the assemblage from the lower Aurignacian (IV) is that
the ornaments seem to cluster in the squares delineated by the possible rock-lined structure
discussed above. Stiner (2010) suggests that many or most of the ornaments may have been
attached to an organic hide or mat, which resulted in dark staining of the sediments observed
during excavation.
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The presence of the rock-lined structure, numerous hearths, and the nature of the
lithic and faunal remains indicates that a range of domestic activities occurred at the site.
This implies that Klissoura Cave 1 was probably a base camp or short-term residential site
during much of its history.
SUMMARY AND GOALS
Considering all of the lines of evidence discussed above, Klissoura Cave 1 is an
extremely important site for understanding behavioral change and shifting foraging strategies
over the course of the Middle Paleolithic through Mesolithic in southern Greece. The length
of the sequence, with the exception of a few hiatuses, and intact stratigraphy fills many gaps
in our understanding of the Greek Paleolithic, in particular the Early Upper Paleolithic
(Uluzzian layer V) and Aurignacian. In addition, advanced excavation methods and
specialized analyses (specifically macrobotanical remains, phytolith analyses, and
micromorphology) provide many avenues of understanding not available to previously
excavated sites. The goals of the larger Klissoura Cave 1 project are to better understand the
Greek Paleolithic through Mesolithic as a whole, examining how stone tool types, use of
ornaments as symbols, and hunting patterns fit with other sites in Greece, and how the site
fills gaps in our understanding of Greece before the Neolithic. Additionally, though it is less
than a day’s walk from the Gulf of Argos, it is the only well-published inland site on
Peloponnese. Thus, intensification and dietary expansion that is recorded at the end of the
Upper Paleolithic at other sites (e.g., Franchthi Cave) may take on a different dimension at
Klissoura Cave 1 in the absence of marine resources.
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The well-preserved faunal remains from the Middle Paleolithic through Mesolithic
layers of Klissoura Cave 1 are the focus of this dissertation. In the coming chapters, I
examine dietary change in the Klissoura Cave 1 sequence from the perspective of animal
exploitation, using zooarchaeological and taphonomic methods and drawing on concepts of
evolutionary ecology discussed in Chapter 1. Changes in environmental conditions are
evaluated (Chapter 3) before examining shifts in prey choice, to distinguish as best as
possible between natural and human processes affecting prey availability. Evidence for nonhuman taphonomic factors in the assemblages is examined (Chapter 4) in addition to
questions about human decisions surrounding the partitioning, transport and butchery of large
and small-bodied prey (Chapters 5 and 6). Spatial variation in modified faunal remains is
also considered within the layers that contain large samples (Chapter 7). The coming chapters
are meant to highlight the uniqueness of Klissoura Cave 1 as a fully terrestrial site on one of
three major southern European peninsulas, while considering the locality as part of a larger
Paleolithic landscape within Greece. Ultimately, the goals of this dissertation are to
understand diachronic trends at a site that spans much of the late Pleistocene and early
Holocene, and use this sequence to augment gaps in our understanding of the evolution of
human subsistence strategies during the Paleolithic in this part of the Mediterranean Basin.
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CHAPTER 3: LATE PLEISTOCENE CLIMATE AND ENVIRONMENT
IN THE EASTERN MEDITERRANEAN
INTRODUCTION
Climatic variability was one of the key pressures acting on human groups worldwide
during the late Pleistocene. Changing climates led to shifts in environmental regimes, which
affected the availability of various plant and animal species as biotic communities expanded
and contracted regionally. Clearly, human groups dealt with such change successfully overall
as populations continued to grow during the Middle and Upper Paleolithic, though there were
declines and extinctions on a local level. This variability would have provided an
environmentally unpredictable backdrop for Neandertal populations in Europe and western
Asia, and for the movement of anatomically modern humans into Europe from the Middle
East after 50,000 years ago (Huntley and Allen 2003).
The occupation of Klissoura Cave 1 spans much of the late Pleistocene, from marine
oxygen isotope stage (MIS) 5a through MIS 1 (about 80,000 until 10,000 BP, see Table 2.1)
(Karkanas, personal communication 4/23/10; Kuhn, et al. 2010). This period is marked by a
high degree of climatic variability with oscillations between glacial and interglacial
conditions, as well as smaller stadials and interstadials (Martinson, et al. 1987). The everchanging climate of this time period witnessed a rapid turnover of plant communities,
ranging from open steppe environments during the glacial periods, to an expansion of forests
during interglacials (Huntley and Allen 2003; Macklin, et al. 2002; Van Andel and Tzedakis
1996), which in turn affected the composition of animal communities. Such changes are
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often reflected by the prey spectra in archaeological sites that correspond with the availability
of local resources (Huntley and Allen 2003).
A central goal of this chapter is to tease out possible evidence for human population
growth and behavior from environmental factors based on data on the hunting patterns of
Pleistocene hominins. As such, some reconstruction of the local and regional environment is
crucial. In this chapter, larger-scale climatic changes that affected the Mediterranean Basin
from MIS stage 5a through 1 are discussed, followed by more localized changes that
occurred in Greece, specifically around Klissoura Cave 1 for the latter part of the occupation
of the site. The general availability of animal resources is predicted in light of changing plant
communities in the area. The environmental reconstructions are based almost exclusively on
sediment cores taken from lakes or the Mediterranean Sea, qualified by data on the
macrobotanical remains from Franchthi Cave (Hansen 1978, 1991) in the southern Argolid
and Klissoura Cave 1 (Albert 2010; Karkanas 2010; Ntinou 2010). In general, nonarchaeological contexts were preferred for environmental reconstructions because they offer
a more complete record without an anthropogenic signature. However, the data from
Klissoura Cave 1 are included because they provide the most local reconstructions available,
and the proximity of Franchthi Cave to Klissoura Cave 1 (about 40 km) justifies the inclusion
of Hansen’s work on plant communities and exploitation. All radiocarbon dates presented
here are calibrated, unless otherwise noted.
ENVIRONMENTAL HISTORY OF THE MEDITERRANEAN: MIS 5a THROUGH 1
Climatic fluctuations in the northern hemisphere during the Pleistocene are primarily
understood from long marine cores from the Indian Ocean (Martinson, et al. 1987) and North
114
Atlantic (Bond, et al. 1993; Bond, et al. 1992; Bond and Lotti 1995), and ice cores from
Greenland (Dansgaard, et al. 1993; Grootes, et al. 1993). The use of these records as a
baseline for other Pleistocene environmental reconstructions is fairly standard (see below).
One of the most significant contributions to come out of this research are the marine oxygen
isotope stages (MIS) that chronicle changing ocean temperatures (Martinson, et al. 1987),
which in turn reflect larger atmospheric changes. Martinson et al. (1987:1) provide a
synthesis of the MIS dating scheme by combining radiocarbon and uranium-thorium dates
with stacked oxygen-isotope chronologies and four different “orbital tuning” approaches,
where they correlate climatic events with orbital forcing. They place the following date
ranges for the MIS stages: 1 (0 – 12,050 BP), 2 (12,050 – 24,110 BP), 3 (24,110 – 58,960
BP), 4 (58,960 – 73,910), 5 (73,910 – 79,250 BP, which marks the beginning of MIS 5a)
(Martinson, et al. 1987:20). Figure 3.1 is adapted from Martinson et al. (1987:19) and shows
the fluctuations of δ18O values; positive values indicate generally warmer ocean conditions,
negative values generally cooler conditions. Aside from the major fluctuations, smaller
events are also noted and dated (Martinson, et al. 1987:20). The case will be made below that
the larger-scale fluctuations are an appropriate baseline for analyzing climatic change in the
Mediterranean Basin, though the smaller fluctuations are less useful for understanding local
climatic shifts.
Establishing a link between marine (Bond, et al. 1993; Bond, et al. 1992; Bond and
Lotti 1995) and the classic Greenland ice core records (Dansgaard, et al. 1993; Grootes, et al.
1993) is important in understanding global climate shifts during the Pleistocene. These
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Figure 3.1 Marine oxygen isotope chronology, adapted from Martinson et al. (1987). Bar at the top
indicates period that were warm (w), cold (c) and mild (m), compared to surrounding periods.
studies typically record fluctuations on the millennial scale; as will be discussed below,
higher resolution is often found in local climate records. Ice cores from Greenland indicate
that climatic instability over the course of the last hundred thousand years was extreme, with
the exception of the comparatively stable Holocene (Dansgaard, et al. 1993:220; Grootes, et
al. 1993:553). In the ice cores, a series of temperature events, called Dansgaard-Oeschger (DO) oscillations were identified (Dansgaard, et al. 1993). Bond et al. (1992:247) examined
marine cores from the North Atlantic and identified a series of six so-called Heinrich events
over the course of the last 70,000 years. These events are marked by drops in sea surface
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temperature and salinity, a decrease in the flux of planktonic foraminifera, and a discharge of
icebergs (Bond, et al. 1992:248). Bond et al. (1993:146) establish a link between the D-O
oscillations and Heinrich events, which punctuate the end of D-O oscillations between
20,000 and 80,000 BP. This confirms the relationship between ice sheet melting and
changing atmospheric and ocean temperatures (Bond, et al. 1993:145), though some
additional ice rafting cycles seem to appear outside of the D-O oscillations (Bond and Lotti
1995).
A few Mediterranean-wide climatic studies are published for the late Pleistocene.
Considering several lines of evidence, including ice cover, sea-level, and pollen data from
ice, marine and pollen cores, Van Andel and Tzedakis (1996) present vegetation maps for
several different stages of the late Pleistocene. They determine that MIS 4 was cold, with an
expansion of open vegetation, whereas MIS 3 was variable with many climatic oscillations
(Van Andel and Tzedakis 1996). Macklin et al. (2002) examine aggradation in alluvial
systems in Greece, Libya and Spain. They determine that alluviation occurred when steppe
vegetation expanded during cool, dry stadials (Macklin, et al. 2002:1634). They found that
MIS 4 was quite cold, with aggradation at sites in their study (Macklin, et al. 2002:1638).
Conversely, MIS stages 3 and 2 are marked by multiple periods of alluviation, indicating a
high degree of climatic variability (Macklin, et al. 2002:1639). Both of these studies offer
fairly coarse reconstructions, though they correspond well with broader trends in late
Pleistocene climates. Harrison and Digerfeldt (1993) compiled lake-level data for the
Mediterranean Basin since the last glacial maximum (LGM), with the assumption that high
lake levels are indicative of wetter climatic conditions, at least on a regional basis. They find
that the last high stand of lakes throughout the region occurred at the LGM (Harrison and
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Digerfeldt 1993:241). Subsequent drying began first in the western Mediterranean at about
16,000-15,000 BP (uncal) and later in the eastern Mediterranean between 13,000 and 11,000
BP (uncal). Conditions became wetter by 12,000 BP (uncal) in the west and gradually
ameliorated in the east until about 6,000 BP (uncal) (Harrison and Digerfeldt 1993:241).
Many studies in the Mediterranean Basin have attempted to link local signatures with
the marine and ice core sequences from the Northern Hemisphere. A 48,000 year core from
the Alborán Sea south of Spain was analyzed and correlated to other Iberian sequences, as
well as North Atlantic ice cores (Cacho, et al. 1999; Fletcher and Sánchez Goñi 2008;
Sánchez Goñi, et al. 2002). Heinrich events 1-5 are evidenced in the sequence by drops in sea
surface temperatures and fluctuations in planktonic foraminifera, similar to those found in the
North Atlantic (Cacho, et al. 1999:703). Dansgaard-Oeschger oscillations are also apparent in
the Spanish record, though they are less intense than the observed Heinrich events (Cacho, et
al. 1999:703). Sanchez Goni and colleagues (Fletcher and Sánchez Goñi 2008; Sánchez
Goñi, et al. 2002:104) correlate vegetation changes and sea surface temperatures from the
Alborán core and determine that D-O stadials were dry and cold, while D-O interstadials
were humid and mild. Heinrich events were the driest and coldest periods. They then relate
the Alborán record to an Atlantic sequence off the coast of Portugal, and find broad
agreement between the two cores in the timing of events, though differences are apparent in
the amount of moisture or temperature gradients at the two sites during various Heinrich
events (Sánchez Goñi, et al. 2002:103).
In Italy, the 102,000 year sequence from Lago Grande di Monticchio was correlated
with the marine sequence, as well as the Greenland ice cores (Allen, et al. 1999; Allen and
Huntley 2000; Allen, et al. 2000; Watts, et al. 1996). The Monticchio record matches the
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marine oxygen isotope stages well, with a difference of less than 2,000 years for the major
boundaries between stages (Allen and Huntley 2000:124; Allen, et al. 2000:106). The
sequence also records many fluctuations found in the North Atlantic, including D-O
oscillations during MIS 2 and 3 (Watts, et al. 1996:150) and Heinrich events 1-6, though
there are some variations. Allen and colleagues (Allen, et al. 1999:742; Allen and Huntley
2000:124) find a divergence between the Monticchio and North Atlantic records before
65,000 BP, with the former displaying a more detailed record with additional fluctuations,
and Watts et al. (1996:151) find that Heinrich events 1-6 in the sequence manifest slightly
later than is indicated in the ice cores. Additionally, some seasonal trends were apparent
during interstadials in the high-resolution Italian record that do not appear in the ice core
records (Allen, et al. 2000:108). The Monticchio record also correlates well with other pollen
cores from southern Europe, including Padul (Spain), Lac du Bouchet (France), Valle di
Castiglione (Italy) and Tenaghi Philippon (Greece), though the Monticchio core has higher
resolution and displays some fluctuations not apparent in the other sequences (Allen, et al.
2000:124).
At Soreq Cave in Israel, changes in the proportion of δ18O and δ13C in speleothems
reflect climatic fluctuations over the last 60,000 years (Bar-Matthews, et al. 1997; BarMatthews, et al. 1999). The authors find cold peaks at 46,000, 25,000, 19,000, 16,600 and
between 13,200 and 11,400 BP, which correspond to Heinrich event 5 (H5), H2, the LGM,
H1 and the Younger Dryas, respectively (Bar-Matthews, et al. 1999:89; Bond, et al. 1992).
Heinrich events 3 and 4 are not represented in the Soreq Cave sequence, and there is a cold
event recorded at 35,000 BP that is not found in the Greenland ice core record (Bar-
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Matthews, et al. 1999:89; Bond, et al. 1992). Warm peaks were found at 54,000 and 36,000
BP.
Climatic reconstructions from the Mediterranean Basin tend to compare quite well
with worldwide MIS shifts (Allen and Huntley 2000; Allen, et al. 2000; Cacho, et al. 1999;
Fletcher and Sánchez Goñi 2008; Sánchez Goñi, et al. 2002). In some cases, individual
Heinrich events and D-O oscillations are also identified (Allen, et al. 1999; Allen and
Huntley 2000; Bar-Matthews, et al. 1997; Bar-Matthews, et al. 1999; Cacho, et al. 1999;
Fletcher and Sánchez Goñi 2008; Sánchez Goñi, et al. 2002; Watts, et al. 1996). However,
many of these events manifest themselves differently in terms of length or severity in the
local records, and in many cases certain Heinrich events are entirely absent from
Mediterranean cores (Allen, et al. 1999; Allen and Huntley 2000; Bar-Matthews, et al. 1997;
Bar-Matthews, et al. 1999; Sánchez Goñi, et al. 2002; Watts, et al. 1996). For this reason, the
general character (warm, mild, cold) of the different MIS stages from 5a onward are included
on the axes of many figures in this dissertation. Though the larger curve fits quite well with
the Greek data discussed below, it must be noted that glacial conditions of MIS 4 may have
begun slightly later and MIS stages 2 and 1 began slightly earlier in Greece than in the Indian
Ocean record (Martinson, et al. 1987, see also Figure 3.2). On the other hand, smaller
Heinrich events and D-O oscillations seem to take on a local character across the
Mediterranean, including Greece (see below). Therefore, this presentation uses the MIS shifts
as a general guide for Mediterranean-wide and Greek climatic changes, but local
environmental studies are used to establish finer-scale shifts.
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Figure 3.2 Simplified schematic of Greek paleoclimate studies mentioned in the text. Dark bars =
warm, wet (forest expansion), grey bars = mild (mixed forest-steppe), white bars = dry, open steppe.
Core locations from studies mentioned in this figure are mapped in figure 3.3. Bottom axis
corresponds to MIS stages in Figure 3.1.
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ENVIRONMENTAL HISTORY OF GREECE: MIS 5a THROUGH 1
Northern and Central Greece
Climatic changes in Greece since MIS 5a generally mirror those from the larger
Mediterranean Basin. In some cases, certain climatic events are expressed differently in the
eastern Mediterranean than to the west (see above). Many environmental reconstructions are
available for Greece, particularly in the northern and central part of the country (see Figure
3.3 for core locations mentioned in this chapter). Of these studies, most focus on the LGM
and later (Kotthoff, et al. 2008; Lawson, et al. 2005; Lawson, et al. 2004; Okuda, et al. 1999;
Turner and Sánchez Goñi 1997; Tzedakis 1993, 1994, 1999; Tzedakis, et al. 2004; Tzedakis,
et al. 2002; Wijmstra 1969; Willis 1997), but some longer cores were analyzed as well
(Tzedakis 1993, 1994, 1999; Tzedakis, et al. 2004; Tzedakis, et al. 2002; Wijmstra 1969). A
synthesis of all of the pollen data from Greece, some taken from pollen cores, others from
archaeological sites, is presented in Figure 3.2. This figure indicates periods of expanded
forest communities with increased moisture or changes in the seasonal availability of
moisture, intermediate conditions with mixed forest-steppe environments, and dry periods of
open steppe vegetation. Changes in the vegetation regimes will be used to interpret the
availability of herbivore species below.
Tzedakis et al. (2002) present pollen data from a core taken from the Ioannina Basin
(core I-284). In general, the area was likely a refugium for forest communities throughout the
Pleistocene, but contractions and expansions of these forests correspond fairly well with
environmental changes found elsewhere in Greece. During MIS 5a and into MIS 4 (between
83,000 and 68,000 BP), forest communities were widespread in the area (Tzedakis, et al.
2002:2044). An earlier long core analyzed by Tzedakis (1993:437; 1994:414), also from the
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Figure 3.3 Map of Greece indicating locations of pollen cores discussed in the text, and in Figure 3.2.
Ioannina Basin (core I-249), indicates that MIS 5a was marked by Quercus (oak) forests with
Ulmus (elm) and Carpinus (hornbeam). These conditions are also found in central Greece,
where a pollen core from Lake Kopais (core K93) has a high incidence of arboreal pollen in
MIS 5a and into MIS 4, from 84,000 to 74,000 BP (Tzedakis 1999:428). A core from the
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Tenaghi Philippon in northeastern Greece also found expanded Quercus ilex (holm oak) and
Pinus cf. nigra (European black pine) forests during MIS 5a (Wijmstra 1969:524).
Forest communities contracted in the Ioannina Basin between 68,000 and 59,000 BP
(MIS 4), and open vegetation with scattered woodlands appeared for the duration of this
period (Tzedakis, et al. 2002:2045). The resolution for the other Ioannina core (Tzedakis
1993, 1994) is exceptionally coarse during MIS 4-2. Transitional parkland, grassland steppe,
and desert steppe communities were identified in this sequence, but no temporal control was
established, so this core will not be included any further in the discussion of these periods.
The study of Lake Kopais also has somewhat lower resolution in MIS 4 between 74,000 and
59,000 BP, but there were probably oscillating conditions with an initial decrease of forest
communities, followed by the expansion of forests, and finally the appearance of more open
vegetation (Tzedakis 1999:431). In the Tenaghi Philippon sequence, the vegetation shifted
from mixed European black pine forest to open Artemisia and Chenopodiaceae steppe during
MIS 4, but there are no dates for this change (Wijmstra 1969:524).
The majority of MIS 3 in the Ioannina Basin between 59,000 and 26,000 BP was
represented by intermediate forest communities, with some short intervals of forest
expansion and contraction (Tzedakis, et al. 2002:2045). Data from the Tenaghi Philippon
sequence, which dates to the latter half of MIS 3, indicates shifts between mixed European
black pine forest and dry, open Artemisia and Chenopodiaceae (Wijmstra 1969:525). This
period in the Lake Kopais core also shows a continuation of oscillating forest and open
conditions, with expansions and contractions of oak, Pinus (pine), Juniperus (juniper) and
Abies (fir) (Tzedakis 1999:431). Lawson et al. (2004) reanalyze the later portion of core I284 from Ioannina and incorporate recent radiocarbon dates into the analysis. They look at
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pollen and microfauna as well as sedimentological studies to reconstruct terminal
Pleistocene/Holocene environments in the region. The earliest part of the sequence dates to
the full glacial, starting at the end of MIS 3, from about 25,150 to 18,650 BP. The pollen is
mostly from herbaceous taxa indicative of an arid environment, and the authors propose that
there was slightly moister Gramineae (grass) dominated steppe interspersed with shrubby
Artemisia and Chenopodiaceae semi-desert (Lawson, et al. 2004:1614). Low frequencies of
oak and pine pollen also persist in the sequence, so there were probably small stands of
deciduous trees in some places on the landscape.
A fairly robust pollen record is available for the late glacial and Holocene, between
26,000 and 11,500 BP, in northern and central Greece. In the Ioannina Basin, more open
vegetation with scattered woodlands appear late in MIS 3 and then persist until the early
Holocene (Tzedakis, et al. 2002:2045). Lawson et al. (2004) finds that the late glacial pollen
record of MIS 2 in the same region, from 18,650 to 11,280 BP, indicates warmer, wetter
conditions than the preceding full glacial. This period is transitional between the open glacial
landscape and interglacial forests, with steppe-grassland and a few trees evidenced by
increasing frequencies of oak, Artemisia and Chenopodiaceae, and corresponding decreases
in pine and Gramineae (Lawson, et al. 2004:1615). The Tenaghi Philippon sequence shows a
gradual amelioration of the climate with an increase in forested conditions, though periods of
open steppe communities continue to be interspersed between mixed or forested conditions
(Wijmstra 1969:525). Lawson et al. (2005) analyze a series of cores from northern Greece,
the longest of which is Nisi Fen, which spans from the late glacial through the Holocene.
They find that from 20,000 to 12,000 BP, the area was mostly open, but had some small
stands of Betula (birch), pine, juniper, and variable amounts of deciduous oak (Lawson, et al.
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2005:883). To the west, Willis (1997) looked at pollen cores from Gramousti Lake and
Rezina Marsh, and Turner and Sánchez Goñi (1997) examined cores from Lake Ziros and
Lake Tseravinas, all of which are in Epirus. Vegetation during the glacial maximum, between
18,350 and 17,900 BP (and possibly earlier), was steppe grassland with Artemisia and
Chenopodiaceae, as well as grasses and herbaceous taxa (Turner and Sánchez Goñi
1997:580). After this there was an expansion of oak woodland, resulting in a mixed foreststeppe from 17,900 to 13,020 BP (Turner and Sánchez Goñi 1997:581). The mixed foreststeppe persisted until 11,405, though pine largely replaced oak in the woodlands (Turner and
Sánchez Goñi 1997:581). During the late glacial, from 15,350 to 11,205 BP, Willis
(1997:404-407) finds open steppe environments, with a slight expansion of oak at about
13,020 in the same region.
Kotthoff et al. (2008) examine pollen from a marine core (SL 152) in the Aegean Sea
that spans the last 20,000 years. During the late glacial, from 20,800 to 14,600 BP, steppe
vegetation dominates the sequence, with Artemisia and Chenopodiaceae persisting, and an
increase of Ephedra (Kotthoff, et al. 2008:1022). Following this, from 14,600 to 12,700 BP,
there is an expansion of broadleaf arboreal vegetation, though steppe communities persist
(Kotthoff, et al. 2008:1025). During the Younger Dryas, from 12,700 to 11,700 BP,
Artemisia becomes more common, followed by increases of Chenopodiaceae and Ephedra,
indicating dry, open conditions (Kotthoff, et al. 2008:1025). In the Lake Kopais core, MIS 2
is represented by full glacial conditions, with high incidences of Gramineae, Artemisia and
Chenopodiaceae pollen and a low frequency of arboreal pollen, indicating dry, open
conditions (Tzedakis 1999:428). Okuda et al. (1999) also analyze a core from Lake Kopais
that encompasses that late Pleistocene and Holocene. In a pollen zone that dates to earlier
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than 14,750 BP (though there is no maximum date), the authors find an open steppe
environment with Chenopodiaceae and Artemisia, which is thought to indicate glacial
conditions (Okuda, et al. 1999:453). From 14,750 to 12,400, there was an expansion of open
oak and pine forests (Okuda, et al. 1999:453), with increases of pine and Ephedra pollen.
In northern and central Greece the Holocene was marked by forest expansion.
Tzedakis et al. (2002:2044) observe forest expansion in the Ioannina Basin between 11,500
and 5,000 BP. Other studies also find forest expansion during the Holocene in this region,
with an increase in oak, Ostrya (hop-hornbeam), and elm pollen, with the addition of Olea
(olive) and fir (Tzedakis 1993, 1994). Lawson et al. (2004:1615) find changes in forest
composition during the early Holocene in the Ioannina Basin, from 11,280 to 6,295 BP.
Deciduous oak forests transitioned to mixed woodlands at this time, with increased quantities
of Ostrya carpinifolia/Carpinus orientalis (European hop-hornbeam/Oriental hornbeam),
Fraxinus ornus (Southern European flowering ash), and C. betulus (European hornbeam).
Wijmstra (1969:525) also found a shift to forests at around 11,600 BP in the Tenaghi
Philippon core. In the early Holocene, from 12,000 to 5,000 BP, the record at Nisi Fen
indicates an expansion of forests, in particular deciduous oak and pine, though other taxa,
including Ostrya-type, Tilia cordata-type (small-leaved lime), European hornbeam, Southern
European flowering ash, and Pistacia (pistachio) also become more common (Lawson, et al.
2005:883). During the transition from the late glacial to the early postglacial from 11,205 to
10,495 BP, Willis (1997:408) finds a decrease in open herbaceous species, such as grasses
and sedges, at the expense of expanding oak and pine forests in the Gramousti and Rezina
cores. Following this, arboreal taxa in the record become increasingly diverse, though a
mosaic of trees, shrubs and herbs persisted between 10,495 and 8,330 BP (Willis 1997:409).
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Turner and Sánchez Goñi (1997:582) also find a continued spread of open forests during the
early Holocene in Epirus. Kotthoff et al. (2008:1025) find increases in non-steppe herb
pollen and a minor increase in non-saccate arboreal pollen (such as oak) starting in the early
Holocene, about 11,700 BP. Precipitation increases after 10,200 BP led to the expansion of
Holocene forests (Kotthoff, et al. 2008:1026). The Holocene in central Greece is also marked
by expanded forests after 12,400 BP, with deciduous oak, Corylus (hazel) and
hornbeam/hop-hornbeam (Okuda, et al. 1999:453).
Southern Greece
The climatic record for southern Greece is sparse compared to the north. No
reconstructions exist for MIS 4 or 5, though there are some fairly complete records after
about 45,000 BP (Albert 2010; Geraga, et al. 2005; Hansen 1978, 1991; Karkanas 2010;
Ntinou 2010). Geraga et al. (2005) identify a series of stadials and interstadials from MIS 3
and later using planktonic foraminifera, pollen, and microfauna δ18O values from a sediment
core taken from the Cretan Basin in the southern Aegean Sea. Pollen indicative of wet,
forested conditions, including Quercus ilex/coccifera (holm/Kemes oak), hazel, hornbeam
and elm is found between 42,000 and 35,000 BP, with a slight contraction of the forests
between 41,000 and 40,000 BP (Geraga, et al. 2005:321) (Figure 3.2). The pollen record is
interrupted between 35,000 and 25,000 BP. From 25,000 to 24,000 the vegetation was
mixed, followed by an expansion of shrubby Artemisia and Ephedra between 24,000 and
11,000 BP, indicative of a steppe environment (Geraga, et al. 2005:321). After 11,000 BP
there was an expansion of forests and modern Mediterranean taxa, such as pistachio, olive
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and Phillyrea, that lasted until 8,000 BP, when a more mixed environment appeared until
7,000 BP (Geraga, et al. 2005:324).
Hansen (1978; 1991) presents the macrobotanical remains from Franchthi Cave in
southeastern Peloponnese (Figure 3.3). Because Franchthi is an archaeological site with
several fairly large depositional hiatuses (see Chapter 2), the data from Hansen (1978; 1991)
are fairly coarse-grained and patchy (Figure 3.2). The earliest archaeological sequence dates
to between 39,280 and 20,000 BP (De Vivo, et al. 2001; Farrand 2000), and is marked by a
cold, dry climate with herbaceous plants, including Lithospermum arvense (corn gromwell,
now called Buglossoides arvensis), Anchusa sp. and Alkanna cf. orientalis, comprising a
steppe environment (Hansen 1991:105). Between 15,000 and 10,500, there was an expansion
of open woodlands, evidenced by the appearance of pistachios, almonds, pears, bitter vetch,
lentils, vetch, wild oats, and wild barley (Hansen 1991:160), as well as arboreal species
indicative of increased moisture and higher temperatures (Hansen 1991:161). Other
macrobotanical data from Franchthi post-date the occupation of Klissoura Cave 1.
A reconstruction of the immediate environment of the Klissoura Gorge (Prosymni)
during the later part of MIS 3 and MIS 2 is possible based on evidence from Klissoura Cave
1. Albert (2010) and Ntinou (2010) analyzed phytoliths and charcoal, respectively, from the
Upper Paleolithic and later components of the site. In general, they found that the local
environment during the Aurignacian occupations of the cave during the later part of MIS 3
was milder than that of the post-LGM Epigravettian (Figure 3.2). No data were available for
the Mesolithic, and preliminary geoarchaeological interpretations of the Middle Paleolithic
simply indicate that the environment was more humid at that time than it was during later
periods (Sitlivy, et al. 2007:4).
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Ntinou (2010) identified plants from the Prunus genus (which includes almonds,
Webb’s almond, and blackthorn) throughout the Upper Paleolithic and Epigravettian layers,
indicating dry, open parkland in the vicinity of the cave. Additionally, during the Early
Upper Paleolithic and Aurignacian, deciduous oak, elm, Acer (maple), and hornbeam/hophornbeam were identified (Ntinou 2010:55). These are mesophilous taxa, but thin growth
rings observed on some of the samples indicate that the trees were probably growing with the
minimum amount of water necessary for survival (Ntinou 2010:56). During the Upper
Paleolithic, Klissoura Gorge was probably a mosaic of open parkland and open woodland,
with Prunus on the hillsides and stands of mesophilous trees on the valley floor (Ntinou
2010:56). A diverse array of thermophilous species, including olive, evergreen oak and
pistachio are found, particularly in Aurignacian layer IV. Ntinou (2010:56) argues that these
warm-loving species are indicative of a refugium in northern Peloponnese during an
interstadial of MIS 3. Together, the mesophilous and thermophilous species point to mild
winter temperatures and just enough precipitation to support the growth of temperate trees
(Ntinou 2010:56).
Based on phytolith analysis, Albert (2010:78) finds an abundance of C3 grasses and
sedges in Aurignacian layer IV, which indicates that the environment was more humid at this
time than in later periods of occupation. The overlying Aurignacian layers (IIIe and IIIg) are
dominated by monocots and a low diversity of C3 grasses, also indicative of greater humidity
(Albert 2010:81). During this period, there was an influx of inflorescence phytoliths,
meaning that grass plants were brought into the site during the flowering season (Albert
2010:81). Other components of the layer III series, III and III’ (which is not actually
considered to be an Aurignacian layer, see Chapter 2), contain C3 grasses and reeds, such as
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Arundo donax (giant cane) (Albert 2010:81). These plants may have grown near the
perennial river that ran through the gorge. C3 grasses are generally indicative of more
temperate environments, and Arundo donax requires fairly moist soils, often growing along
rivers and in marshy areas (Turland, et al. 1993:164; Tutin, et al. 1980:253). These findings
support Ntinou’s (2010) assessment that the Upper Paleolithic climate in the vicinity of
Klissoura Gorge was fairly mild. The only phytoliths found in Layer 6 are from C3 grasses,
but these samples come from a disturbed area designated as a pit so it is unclear if they are
associated with the Upper Paleolithic or later periods.
As discussed above, environmental conditions changed throughout the Mediterranean
during the LGM, but this period is not represented in the Klissoura Cave 1 sequence
(Karkanas 2010; Ntinou 2010). The LGM is also missing from Franchthi Cave (Farrand
2000; Hansen 1991), so this may represent a depositional hiatus in southern Greece, and
possibly also a movement of human populations out of the region at this time. After the
LGM, the vegetation of the gorge reflects a drying trend and a more open environment
(Albert 2010; Ntinou 2010). Prunus remained dominant in the Epigravettian layers (Ntinou
2010:55). Phytoliths from C4 grasses, which are indicative of a drier, warmer environment,
are found in addition to C3 grasses and reeds in the Epigravettian layers (Albert 2010:82).
These lines of evidence point to dry, open parkland in the area around Klissoura Cave 1 in
this later period, in contrast to a more temperate Upper Paleolithic environment (Albert 2010;
Ntinou 2010).
Geoarchaeological lines of evidence offer a fairly low-resolution picture of climatic
conditions during the Klissoura Cave 1 occupation. Karkanas (2010:32) notes that the
uppermost layers (Mesolithic and Epigravettian) have more in common depositionally with
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the late Middle Paleolithic than with the intervening Aurignacian. This is because
sedimentation during the Aurignacian can primarily be attributed to anthropogenic processes,
whereas natural processes, particularly the introduction of clastic sediments into the site,
caused the majority of sedimentation in the earlier and later periods (Karkanas 2010:32).
Based on this, Karkanas (2010:32) suggests two possible scenarios for the accumulation of
clastic materials at Klissoura Cave 1 during the Mesolithic and Epigravettian: either
precipitation rates were higher following the LGM, which caused more runoff and slope
wash into the cave, or a cooling and drying trend led to a more open environment in the
gorge, leaving the hillsides destabilized and more vulnerable to erosion during occasional
storms. Karkanas (2010) points out that the latter scenario is supported by the botanical
results of Albert (2010) and Ntinou (2010). The older Upper Paleolithic layers, which include
from top to bottom a Mediterranean backed bladelet industry (III’), another non-Aurignacian
Upper Paleolithic industry (III”), and multiple Aurignacian layers, have loose, dusty-looking
sediments that lack recrystallized or cemented wood ash, indicating that they formed under
periods of relatively low humidity (Karkanas 2010:33), though this may reflect conditions
inside of the shelter. Despite decreased humidity at this time, there was clearly enough
moisture to sustain temperate tree populations in the valleys (Ntinou 2010) and a more stable
landscape (Karkanas 2010:34).
To date, botanical, phytolith and geoarchaeological analyses of the Middle Paleolithic
at Klissoura Cave 1 are ongoing. General observations of the stratigraphy and formation of
the Middle Paleolithic sediments indicate that they occurred under more humid conditions
than those in the Upper Paleolithic (Sitlivy, et al. 2007:4). Karkanas (2010:32) notes that the
late Middle Paleolithic layers may have formed under similar conditions to the Mesolithic
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and Epigravettian, so as with the later layers, it is unclear if during the Middle Paleolithic the
environment was wetter, causing an influx of clastic sediments, or if it was cooler and drier,
with a decrease in vegetation and subsequent destabilization of the landscape. Once any
botanical and phytolith samples from the Middle Paleolithic are analyzed, this picture will
hopefully become clearer. At this point, however, we must rely on non-local climatic
reconstructions from northern Greece to try to understand possible changes in the
composition of faunas in the Middle Paleolithic layers.
Environmental Conditions during the Occupation of Klissoura Cave 1
The end of MIS 5 is represented consistently in northern and central Greece by
expanded forest conditions (Tzedakis 1993, 1994, 1999; Tzedakis, et al. 2002; Wijmstra
1969). The earliest part of the archaeological record at Klissoura Cave 1 (Middle Paleolithic
XVIII-XXb) was probably deposited at this time, based on the lithic and geoarchaeological
data (Karkanas, personal communication 4/23/10). However, no dates are currently available
for the lowest layers of Klissoura Cave 1, so this assumption may be revised in the future. An
alternative possibility is that the Middle Paleolithic XX series dates to MIS 5a, and Middle
Paleolithic layers XVIII-XIX were deposited during MIS 4 (Karkanas, personal
communication 4/23/10). This means that the earliest layers either formed during the milder
forested or mixed forested conditions of MIS 5a, or during the open steppe conditions of MIS
4 (Tzedakis 1993, 1994, 1999; Tzedakis, et al. 2002). For the purposes of this discussion, in
keeping with the lithics data, we will assume that Middle Paleolithic layers XVIII-XXb were
either deposited in MIS 5a or 4, and overlying undated Middle Paleolithic layers (X-XVII)
formed during MIS 4 or 3 (Table 3.1).
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Table 3.1 Taxa expected in the vicinity of Klissoura Cave 1 by MIS stage. Note that the MIS stage of
the lower layers are unknown, which is indicated by "/" and both options are included. MIS 3 is
highly variable and local reconstructions are available for the UP layers but not the MP. P = present,
A = absent, ? = unsure.
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In general, environmental data from southern Greece correspond fairly well with
those from northern and central Greece for the latter half of MIS 3 and later periods, though
the records are quite incomplete. Small disagreements among records likely reflect local
conditions, as some areas seem to lag behind others in vegetation and climatic changes
(Figure 3.2). The general picture for MIS 3 in Greece seems to be a high degree of climatic
variability, but with generally more amicable conditions. Both continental and southern
Greece oscillated between mixed forest, steppe communities and warm, forested conditions,
with just a few short periods of drying (Albert 2010; Geraga, et al. 2005; Karkanas 2010;
Ntinou 2010; Tzedakis 1999; Tzedakis, et al. 2002; Wijmstra 1969) (Figure 3.2). These were
the conditions during the later Middle Paleolithic occupations (VIII and possibly X-XVII) of
Klissoura Cave 1, as well as the earliest Upper Paleolithic, with the Early Upper Paleolithic
(Uluzzian layer V), Aurignacian series (IV, IIIe-g, IIIb-d), Upper Paleolithic (nonAurignacian) industry (III”) and Mediterranean backed-blade industry (III’) deposited during
this time. Unfortunately, the limited resolution of the environmental reconstructions coupled
with radiocarbon date error ranges from this period makes it impossible to correlate specific
environmental fluctuations with these various cultural designations. Toward the end of MIS 3
conditions became drier and steppe communities expanded across Greece, which persisted
through the LGM (Geraga, et al. 2005; Hansen 1991; Kotthoff, et al. 2008; Lawson, et al.
2004; Okuda, et al. 1999; Turner and Sánchez Goñi 1997; Tzedakis 1999; Tzedakis, et al.
2002) (Figure 3.2). The LGM is not represented at Klissoura Cave 1 (Albert 2010; Karkanas
2010; Ntinou 2010).
After the LGM, particularly in northern Greece, conditions ameliorated and mixed
forest-steppe communities became widespread (Lawson, et al. 2005; Lawson, et al. 2004;
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Turner and Sánchez Goñi 1997; Wijmstra 1969). In southern Greece, it seems that conditions
remained dry until shortly after 15,000 BP (Albert 2010; Geraga, et al. 2005; Karkanas 2010;
Kotthoff, et al. 2008; Ntinou 2010); this also occurred in some parts of central Greece
(Okuda, et al. 1999; Tzedakis 1999; Tzedakis, et al. 2002; Willis 1997) (Figure 3.2). At
Klissoura Cave 1, the Epigravettian (IIa-d) was deposited during this period of continued dry,
open environments. At the beginning of the Holocene, forests expanded across Greece and
the modern Mediterranean-type plant taxa appeared (Geraga, et al. 2005; Hughes, et al. 2006;
Kotthoff, et al. 2008; Lawson, et al. 2005; Lawson, et al. 2004; Okuda, et al. 1999; Turner
and Sánchez Goñi 1997; Tzedakis 1993, 1994; Tzedakis, et al. 2002; Willis 1997) (Figure
3.2). No botanical or phytolith samples are available for Klissoura Cave 1 for the Mesolithic
(3-5a), though based on widespread climatic amelioration in the rest of Greece, it is likely
that conditions also became more mild in the Argolid.
BIOTIC COMMUNITIES AND RESOURCE AVAILABILITY
Understanding the habitat requirements of prey species at Klissoura Cave 1 is
important in interpreting the suite of taxa hunted by the site occupants at different times
during the Pleistocene. Ultimately, this will aid in determining if changes in species
composition at the site were due to environmental causes or human hunting pressures.
Ungulates and small game are particularly significant, as carnivore species are fairly rare in
the assemblages (see Chapter 5). Large game species found in the assemblages include
European fallow deer (Dama dama), ibex (Capra cf. ibex), European wild ass (Equus
hydruntinus), aurochs (Bos primigenius), wild pig (Sus scrofa), red deer (Cervus elaphus),
roe deer (Capreolus capreolus), and chamois (Rupicapra rupicapra). These taxa can be
136
broadly categorized as grazers and browsers, though considerable flexibility exists in their
diets. Grazers tend to consume more grasses and sedges, while the diets of browsers are
composed of more leaves and vegetation of woody plants and forbs (Janis 2008; Searle and
Shipley 2008). It might be expected, particularly with taxa that adhere more strictly to these
categories, that grazers would be more successful in dry, open habitats, while browsers prefer
wetter, forested environments. Either group should be tolerant of mixed vegetation.
European fallow deer are intermediate feeders. Grasses compose the bulk of their
diets, though they also include herbs, broad-leaf tree browse, shrubs, mast, and forbs on a
seasonal basis (Feldhamer, et al. 1988:317; Nowack 1999:1099). They inhabit a range of
environments, including woodlands, mixed and broad-leafed forests, grasslands, scrub,
savanna, and areas with subalpine vegetation (Feldhamer, et al. 1988:317). Red deer
similarly live in a variety of habitats, including plains and mountainous areas, though they
prefer to have access to forested areas, stands of shrubs, tall grasses or reed thickets for
shelter (Baskin and Danell 2003:56). They subsist on a broader range of plants than most
cervids, feeding primarily on browse, in addition to grasses and herbs (Baskin and Danell
2003:59-61). Their diets are largely seasonal, with grasses, rushes and sedges, heaths, forbs,
deciduous browse and conifers common in the summer, and heaths consumed in the winter
(Latham, et al. 1999:412). There may also be considerable geographical variation in red deer
diets (Baskin and Danell 2003; Latham, et al. 1999), although their ecological niche is wide
in most regions. Roe deer prefer woody and mosaic landscapes with dry plains or steppe and
adjacent small stands of forest, shrubs or grass thickets, though they avoid dense forests
(Baskin and Danell 2003:110). Their diets consist primarily of herbaceous plants, such as
forbs, and to a lesser extent grasses, sedges, rushes, heaths, deciduous browse, conifers and
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ferns; the majority of the latter is consumed during the summer (Baskin and Danell
2003:113; Latham, et al. 1999:412).
Ibex are flexible grazers that subsist primarily on grasses in the spring and summer,
but will also browse on forbs, leaves, shoots and bark during the winter, though they
typically avoid dense forests (Nowack 1999:1223; Roberts 1977:194; Schaller 1977). Ibex
are usually associated with high-altitude environments (Nowack 1999:1223; Roberts
1977:193; Schaller 1977), but Phoca-Cosmetatou (2004b) has argued that this may be a
product of recent marginalization by habitat depletion. Today, in protected areas, ibex
occupy a wide range of ecozones, preferring those that are rugged but not necessarily highaltitude. Chamois are mixed feeders that inhabit alpine meadows during the summer and
spring, moving to lower elevations in the fall and winter (Nowack 1999:1213), where they
occupy rocky mountain forests or cliffs near the edges of forests (Baskin and Danell
2003:312). In the summer they subsist on herbs, grasses and flowers, and in the winter they
consume dry grass, twigs of shrubs, young pine shoots, moss, and lichens (Baskin and Danell
2003:314; Nowack 1999:1213).
It is more difficult to understand the ecology of the European wild ass, Equus
hydruntinus, because the species is extinct. Recent genetic studies have shown that E.
hydruntinus is closely related to Equus hemionus, or the Asian wild ass (Orlando, et al.
2006:2087), so it may be appropriate to use Asian wild ass ecology as a proxy for that of the
European wild ass. It should be noted that samples of European wild ass specimens tested for
genetic material by Orlando et al. (2006:2084) came from areas that were cold steppe and
open, dry grassland at the time that E. hydruntinus inhabited the region. Today, the Asian
wild ass lives in semi-desert and steppe environments (Baskin and Danell 2003:5). They
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subsist mostly in cereal grasses, wormgrass and sedges, though they also consume
herbaceous plants and shrubs to a lesser extent (Baskin and Danell 2003:6). Aurochs, the
wild progenitors to cattle, are also extinct, though several studies were conducted in an
attempt to understand their ecology. Based on their distribution at archaeological and
paleontological sites, climatic reconstructions at these localities indicate that they tended to
live in low-lying, flat floodplain ecosystems (Hall 2008:189; van Vuure 2005:175). Like
modern cattle, and based on morphological observations and early documentation of the
animals before they were extinct, aurochs were likely grazers that mostly ate grasses, sedges
and rushes (van Vuure 2005:213).
Wild pigs have different feeding habits than most other ungulates. They are
omnivores, and feed on a variety of invertebrates, small mammals, bird eggs and carrion, as
well as tree bark, grasses, seeds and acorns (Baskin and Danell 2003:20; Graves 1984:484).
Though they are extremely flexible feeders and are able to tolerate a wide range of habitats,
they require dense stands of forest, shrubs or grass for shelter, typically near water holes,
rivers or swamps (Baskin and Danell 2003:16; Graves 1984:487).
Fallow deer are the most adaptable of the ungulate species found at Klissoura Cave 1,
though red deer, roe deer, chamois and ibex are all comfortable with mixed forest/steppe
ecozones. Ibex are also found in more open environments, which is probably where
European wild ass and aurochs would have occurred as well. Only pigs prefer more forested
areas.
Small game species found in the Klissoura Cave 1 assemblages includes one of three
species of similarly-sized tortoises (Testudo graeca, T. hermanni, or T. marginata), European
hare (Lepus europaeus), rock partridge (Alectoris graeca) and great bustard (Otis tarda).
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Tortoises are fairly flexible in terms of habitat requirements, occupying lush meadows, scrub
covered rocky hillsides, woodlands, and stabilized dunes, though they prefer moderately
dense vegetation (Arnold and Burton 1992). Hares also live in a wide range of environments,
including open fields, grasslands, steppes and woodlands, though they prefer flatter areas
(Burton 1991; Mitchell-Jones, et al. 1999). Rock partridges occupy areas with oak, pine and
juniper, and open stands of beech, hornbeam, oak and oriental plain, preferring very open and
rocky areas near water when nesting (Cramp 1980:659; Handrinos and Akriotis 1997;
Vavalekas, et al. 1993). Great bustards are large ground-dwelling birds that typically inhabit
more open areas, including steppe environments, lowlands, river valleys and treeless plains
(Cramp 1980:659; Heinzel, et al. 1992:128; Johnsgard 1991). Today they are also found in
stubble fields (Lane, et al. 2001).
In general, the important small game species found at Klissoura Cave 1 tend to be
rather non-selective in their habitat requirements. In particular, tortoises, hares and partridges
probably were not affected by century or millennial-scale shifts in plant communities in
southern Greece, because these changes were never that extreme in this part of the
Mediterranean. Great bustards are a notable exception that likely were affected by vegetation
changes. They exhibit a clear preference to open areas; with wingspans up to 2.6 m (Cramp
1980:659), they require treeless environments to effectively take flight.
CONCLUSIONS
In general, changes in the proportions of large game species at Klissoura Cave 1 can
serve as a rough indicator of shifts in plant communities in southern Greece during the
occupation of the site. This of course does not take into account the distances that hominins
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were willing to travel to find game, but this will be addressed in Chapter 6, and was probably
not a major factor. A summary of game animals expected in different archaeological layers,
based on habitat preferences and environmental reconstructions is presented in Table 3.1. In
terms of large game, fallow deer were likely the dominant ungulate species in the vicinity of
Klissoura Cave 1 throughout the Paleolithic and early Holocene sequence. Certain taxa,
including red deer, roe deer and chamois were probably available to hominins occupying the
site when there were mixed open and forested communities in the area. Ibex were present in
periods of mixed steppe/forest environments, or when it was more open. During periods with
dry, open steppe conditions, wild ass and aurochs were more likely to have been an option
for human hunters. Wild pigs were probably only available when conditions were wetter and
there was an expansion of forests.
Based on their habitat requirements, it is likely that small game animals such as
tortoises, hares and partridges were available to humans occupying Klissoura Cave 1
throughout the late Pleistocene. These taxa are found in almost every level of the site,
indicating that they were present in the environment to some degree (see Chapter 5). There is
little reason to believe that environmental changes discussed above would have vastly
changed the frequency of these animals in and around Klissoura Gorge, so their varying
proportions in the archaeological assemblages likely reflects shifts in human hunting
strategies. Conversely, great bustards are sensitive to vegetation shifts, so their presence and
absence likely tracks expanding and contracting grasslands.
Considering only environmental conditions, and depending on the extent of increased
moisture and expanded forests in southern Greece, or if the earliest archaeological layers
were deposited during MIS 5a or 4, two possible suites of prey species were probably
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available during the earliest Middle Paleolithic occupation (XXa-XXb, XVIII-XIX). Fallow
deer and wild pigs, as well as tortoises, hares, and partridges are expected in the vicinity of
Klissoura Cave 1 if these layers were deposited during MIS 5a. Conversely, if the layers
formed during MIS 4, fallow deer, ibex, wild ass, aurochs, and the full suite of small game
(including great bustard), are expected. The situation is similar for the overlying Middle
Paleolithic layers (X-XVII), if they were deposited during MIS 4 or 3. In these instances,
changes in the suite of species found at Klissoura Cave 1 may provide insight into the
specific environmental shifts occurring in southern Greece during the Middle Paleolithic.
This would be apparent in the relative evenness of ungulate species exploited, as well as the
presence or absence of key ungulate species such as wild pig, or wild ass and aurochs.
Changes in the representation of tortoises, hares and partridges would likely have more to do
with human hunting pressures than environmental variability (see Chapter 5), though the
presence of great bustard would probably correlate with an open environment.
Radiocarbon dates indicate that the latest Middle Paleolithic layer (VIII), Early Upper
Paleolithic (Uluzzian layer V), Aurignacian series (IV, IIIe-g, IIIb-d), Upper Paleolithic
(non-Aurignacian) layer III”, and the Mediterranean backed bladelet industry (III’) formed
during MIS 3. The climate was highly variable during this interval and oscillated between
wet, forested conditions, milder phases of mixed forest-steppe vegetation, and dry, open
periods. Mixed forested and opened areas probably persisted in the direct vicinity of
Klissoura Cave 1 at this time, though there was slightly more moisture when layer IV was
deposited. Environmental fluctuations during this period may again be indicated by the
presence or absence of certain ungulate species, or great bustard.
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The Epigravettian (IIa-d) occupation of Klissoura Cave 1 occurred at a time when
dry, open vegetation persisted. It is expected that the full range of small game animals,
including great bustards, were present in the environment, in addition to a fairly limited range
of ungulate species that include fallow deer, ibex, European wild ass, and aurochs. The
Mesolithic (3-5a) that followed was marked by increasing moisture and expanded forests of
the Holocene. Faunal resources available in and around Klissoura Gorge would have
included wild pig and fallow deer, in addition to small game animals such as tortoises, hares
and partridges.
In the chapters that follow, game use and prey selection will be examined from the
Middle Paleolithic through Mesolithic at Klissoura Cave 1. Based on the environmental
reconstructions, habitat requirements of prey species, and predictions discussed above, I will
make the case that some changes in game use across the Pleistocene and early Holocene at
Klissoura Cave 1 were linked to shifts in vegetation communities in southern Greece, while
others were a product of hunting pressures on certain species exploited by hunters in the
region.
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CHAPTER 4: TAPHONOMIC PROCESSES BONE ACCUMULATION AND ATTRITION
INTRODUCTION
A key issue in evaluating zooarchaeological assemblages is determining whether
faunas were accumulated and modified by human foragers, and the extent to which nonhuman processes impacted the remains. At an archaeological site such as Klissoura Cave 1,
with unquestionable human activity throughout the sequence evidenced by an abundance of
stone tools (Kaczanowska, et al. 2010; Koumouzelis, Ginter, et al. 2001; Sitlivy, et al. 2007)
and hearths (Karkanas, et al. 2004; Pawlikowski, et al. 2000), it seems likely that humans
were mostly responsible for the collection of the faunal materials. However, there is no
guarantee that all animals were introduced into the site by humans. Nor is it certain that nonhuman processes, such as weathering, carnivore damage, or in situ attrition, played no role in
shaping the assemblages. Several analyses will be presented here to determine the extent to
which, if any, non-human taphonomic agents impacted the faunas of Klissoura Cave 1. These
include an evaluation of the input of faunal remains to better understand differences in
sample size and faunal density, observations of macroscopic damage caused by weathering,
carnivores and rodents, an analysis of in situ density-mediated attrition, and the degree of
fragmentation of the assemblages to evaluate potential changes in breakage related to time
period or prey body class.
Taphonomic processes are those that act on an animal’s remains after it dies and
include the pre-burial and post-burial histories of the carcass (Lyman 1994:3, see also
Gifford 1981). Taphonomic agents unrelated to human activities include transport and
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gnawing by carnivores (Hockett 1999; Lloveras, et al. 2008a; Stiner 1991, 1994), digestion
(Andrews 1990; Lloveras, et al. 2008a, 2008b, 2008c; Rensberger and Krentz 1988),
chemical and mechanical weathering (Behrensmeyer 1978; Lyman and Fox 1989; PhocaCosmetatou 2005), soil corrosion and diagenesis (Karkanas, et al. 1999; Nielsen-Marsh and
Hedges 2000; Stiner, et al. 2001; Weiner, et al. 1993), rodent gnawing (Brain 1980, 1981;
Hockett 1989; Hoffman and Hays 1987; Maguire, et al. 1980), and root etching (Denys 2002;
Fernandez-Jalvo, et al. 2002; Fisher 1995). Other taphonomic processes are the result of
human hunting and processing behaviors, including burning, carcass transport, trampling,
butchery, bone marrow extraction, and heat-in-liquid grease rendering (Bar-Oz and Munro
2005; Bunn, et al. 1988; Costamagno, et al. 2005; Denys 2002; Fisher 1995; O'Connell, et al.
1988; Outram 2001). Though human processes could easily be included in this chapter, they
are explored separately in Chapter 6, for organizational purposes.
In an archaeological assemblage, there are three overarching processes that affect
faunal remains: accumulation, modification and attrition. Accumulative agents may be
human hunters carrying prey to a camp site, carnivores that bring a kill back to a shelter, or a
fluvial system that washes objects to a particular area. Modifications include processes that
may affect the appearance of bone while not always causing bone loss, such as low levels of
carnivore and rodent gnawing, weathering, and human butchering or burning. Agents that
cause attrition, or bone loss, include chemical or mechanical processes that lead to bone
diagenesis, carnivore gnawing and digestion, and human processing. In some cases, those
agents responsible for accumulation or modification may also cause attrition. For example,
carnivores can bring kills to a rock shelter or cave, but they can also move materials away
from an archaeological site during scavenging or destroy bones entirely. Aside from human
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subsistence behaviors such as the selective accumulation of meaty body parts or high-ranked
species, burning, and butchery damage (see Chapters 5 and 6), the main non-human
taphonomic processes that may have affected the Klissoura Cave 1 faunas are weathering and
diagenesis, trampling, bioturbation, and carnivore accumulation and gnawing.
METHODS
A macroscopic survey of the Klissoura Cave 1 faunas indicate that overall there is
good preservation of the materials. Excavation recovery methods included screening and the
recovery of both shaft and spongy bone fragments. Elements representing a range of
structural densities (see Lyman 1984; 1994) were recovered, in addition to fragile fetal
elements in many layers (Chapter 6). Variation in bone condition exists through the
sequence; the oldest Middle Paleolithic assemblages (XI-XIV and lower) are highly
mineralized and some display a metallic blue patina on their surfaces (Figure 4.1). Most
Upper Paleolithic remains are covered by a layer of solidified ash that could not be removed
without damaging the surface of the bones (Figure 4.2). Attempts to remove the ash on bird
bones using acetic acid proved to be ineffective and potentially damaging to the materials
(Bocheński and Tomek 2010). This coating likely obscures some of the butchery damage, in
particular fine tool marks, though impact fractures and green bone breaks were still easily
discernable. Because bone surfaces in the Upper Paleolithic assemblages were often
obscured, there are some limitations when it comes to discussing cut marks, which will be
addressed in Chapter 6. The skeletal remains are highly fragmented, particularly in the
Middle Paleolithic layers, and the only intact elements are small, compact tarsal and carpal
bones and a few phalanges. Excavation damage is uncommon in the assemblages, and is
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obvious when it occurs because the broken edges are noticeably lighter than the rest of the
bone. Though the materials are highly fragmented, many of the specimens are easily
identified to species or body size class and to skeletal element, based on diagnostic features
or “landmarks.”
Figure 4.1 Metallic blue patina on Middle Paleolithic long bone fragment. From layer XIX.
Bioturbation does not seem to be a major factor at the site. A few micromammals
were observed in the assemblages, but their frequencies were quite low and rodent damage is
uncommon (see below). In general the stratigraphy is fairly intact and was analyzed carefully
(Karkanas 2010). Areas of the site thought to be intrusive pits or mixed during excavation or
geoarchaeological analysis were excluded from the zooarchaeological analysis. There is a
slight amount of mixing between the earliest Upper Paleolithic and latest Middle Paleolithic
layers, evidenced by the presence of a few shell ornaments in the Middle Paleolithic layers
VI and VII in the area immediately below the Aurignacian shelter feature (Stiner 2010), so
layers VI and VII are excluded from the analysis. The ornaments are small in size (mean 1.28
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cm), and could have moved between layers when humans disturbed the sediments or fallen
into rodent burrows. The lithic industries in Layer VIII and below are Middle Paleolithic.
Figure 4.2 (a) Light carbonate fraction on great bustard tibiotarsus. From disturbed zone (layer 6a).
(b) Heavy carbonate fraction on great bustard lumbar vertebrae. From disturbed zone (layer 6).
The analytical units and techniques for the zooarchaeological analyses follow those
outlined in Grayson (1984), Klein and Cruz-Uribe (1984), Stiner (1994) and Lyman (1994).
The Wiener Laboratory faunal reference collection at the American School of Classical
Studies at Athens and unpublished electronic faunal manuals created by Mary Stiner were
used to identify the materials. The basic counting unit in the analysis is number of identified
specimens or NISP (Grayson 1984). The term as used here refers to every fragment or whole
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bone that could be identified to element and body class, or more specific designation.
Specimens were identified to the greatest precision possible: genus (e.g., Testudo sp.),
species (e.g., Dama dama) or body size category (e.g., medium ungulate or small mammal).
In the case of articulated elements or mandible portions with teeth, each individual element
that could potentially be separated was counted individually to control for variation in postdepositional processes (Stiner 1994) but with articulations noted. For example, the NISP of a
complete right mandible of an adult fallow deer with all of the teeth intact would be eleven.
The counting unit NISP is fundamental but is not sufficient for all analyses. It does not, for
example, take into account fragmentation or identifiability that may differ among species, or
differential preservation (Grayson 1984:21-23; Lyman 2008:29-30). Because of this, some
derived units are used in addition to NISP.
Minimum number of elements, or MNE, is derived from NISP by recording unique
features on each element and tabulating the number of times each landmark occurs in an
assemblage for a species or ungulate body size group. For example, if there are four medial
distal femur fragments from Dama dama in a given layer, the MNE is four. Side is not taken
into account when calculating MNE. In the case of long bones, landmarks on both epiphyses
(ends) and diaphyses (shafts) are considered. Another commonly used counting unit is
minimum number of individuals, or MNI, which is the smallest number of animals necessary
to account for the faunal remains at a site (Grayson 1984:27). It is derived from the highest
count of the most commonly occurring element (MNE) in an assemblage, divided by the
number of times that particular element occurs in the body. MNI is used less in this study
than NISP and MNE, for reasons that will be explained in Chapter 5.
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Other observations were recorded for each specimen (following Stiner 1994, 2005),
including fusion state in the case of bones, wear stages for mandibular teeth, body side,
burning damage and intensity (Stiner, et al. 1995), and surface damage from weathering,
plant roots, gnawing rodents and carnivores, and tool marks. The coding of elements,
portion-of-element, age criteria, and taphonomic variables follows Stiner (1994; 2005). The
maximum length of each specimen was recorded to control for the relationship between
fragmentation and differences in prey body size. It must be noted that sample totals in
various tables in this dissertation vary according to whether the analysis is confined to
specific-specific data or includes specimens that could be identified only to body size class.
Shattered dental specimens or overly-recognizable bone fragments (such as tiny pieces of
tortoise shell or the anterior groove on cervid metatarsals) pose similar problems and are
excluded from tables that indicate species frequencies. As an example, Table 4.1 illustrates
the overrepresentation of cervids that would result from including the anterior groove of
metatarsals. These specimens are included in the taphonomic assessments, however, such as
overall degree of burning or fragmentation rates.
Faunal assemblages in the different layers at Klissoura Cave 1 vary in size, though
not to a great degree (Table 4.2). Some parts of the uppermost layers were examined by
previous studies (Koumouzelis, Ginter, et al. 2001; Koumouzelis, Kozlowski, et al. 2001;
Tomek and Bocheński 2002) and were unavailable for analysis. In addition, the excavated
area of the site was not the same for the entire sequence because the excavation grid was
reduced downwardly from 14 m2 in the uppermost layers to 6 m2 in the lowest layers (see
Chapter 7). Lateral variation in material density in the sediments is also noted in layers
representing the most intensive occupations or those that experienced less erosion. Finally, a
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Table 4.1 Fallow deer metatarsal NISP counts and NISP:MNE ratios that include anterior groove
fragments and corrected counts that exclude anterior groove fragments, compared to metacarpal NISP
counts and ratios. Metacarpals were used for comparison because their structural density is similar
and they are equally recognizable in an assemblage as are metatarsals. NISP counts for metatarsals
exclude anterior grooves for taxonomic frequencies because they over-represent fallow deer
frequencies. Only Middle Paleolithic layers are depicted because they are the most extreme, but
cervid metatarsal anterior grooves (and other overly-recognizable fragments such as unidentifiable
tooth and tortoise shell fragments) are excluded in all layers.
small portion of the abundant materials from the Aurignacian series IIIe-g and nonAurignacian Upper Paleolithic (III”) were excluded from the sample due to time constraints,
though the majority of the bones from these layers were analyzed. An analysis of the input of
faunal materials relative to sediment volume in the different layers of Klissoura Cave 1 was
conducted following Stiner and Munro (2011). This was done in order to understand if
sample size differences between the assemblages can be explained by differences in
excavated volumes, and the relationship between the accumulation of sediments, artifacts,
and faunal remains in each layer. Faunal input (or density) was evaluated by dividing
vertebrate NISP counts by the sediment volume in liters for each archaeological layer, or
input = NISP/L. Some attempt can be made to correct accumulation rates by dividing input
by the time interval represented by depositional layers given a reliable set of radiometric
dates, which are lacking in the Middle Paleolithic layers of the site. Accumulation rates for
the faunal remains can then be compared to our understanding of the site from a
geoarchaeological standpoint. According to Karkanas (2010) the sediments of the earlier
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Upper Paleolithic have a strong anthropogenic signature, while the Middle Paleolithic and
post-LGM layers have a greater input of geogenic sediments.
Table 4.2 Rates of faunal input (or faunal density) by layer.
Two methods were used to determine rate of bone fragmentation for different layers
at the site. Maximum length was recorded for all specimens, assuming they were not broken
during excavation. Specimens were then grouped by body class for each layer to correct for
structural differences between prey of different sizes and locomotion, for example ungulates
and birds. The means for each group were compared by layer to determine if there was a
change in fragmentation through the sequence that may reflect intentional bone breakage or
post-depositional processes such as trampling. The second method used to examine
fragmentation rates between levels is the index MNE/NISP, which provides a standardized
value between 0 and 1, where smaller numbers reflect greater rates of fragmentation. The
MNE variable is correlated to NISP but the index is valuable for isolating differences in the
degree of fragmentation within a prey size class. In examining the fragmentation index, only
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medium ungulate remains were included because they (mostly fallow deer) represent the bulk
of the assemblage throughout the sequence (see Chapter 5).
Klissoura Cave 1 is a rock shelter, which makes it an attractive refuge for hominins as
well as carnivores. Like humans, large carnivores have the potential to accumulate the
remains of large-bodied prey at a site. Small and large carnivores can also impact
archaeological faunas by denning, which can mix sediments, and gnawing on human food
refuse. Many mammalian carnivores are recorded in Greece during the Late Pleistocene,
including Ursus arctos (brown bear), Ursus spelaeus (cave bear), Canis lupus (wolf), Vulpes
vulpes (red fox), Crocuta sp. (hyena), Panthera sp. (lion, leopard), Lynx lynx (Eurasian lynx),
Felis silvestris (wild cat), and various mustelids such as martins (Martes sp.) and badger
(Meles meles) (Tsoukala 1992). Mammalian carnivore influences on an archaeological
assemblages can be assessed through several lines of evidence: the presence of carnivore
bones that lack human modifications, prey body part profiles that match known carnivore
transport preferences (Stiner 1991), fetal or neonate carnivore remains that indicate the
shelter was used as a den (Stiner 1994), coprolites, and carnivore damage on the bones
themselves (Maguire, et al. 1980; Marean and Spencer 1991; Stiner 1994). Stiner (1991;
1994:249-258) argued that assemblages accumulated by Paleolithic hominins tend to be more
complete in their representation of ungulate body parts (with the exception of some headdominated Middle Paleolithic faunas), whereas canid and hyena accumulations often contain
more head parts, or feet. Ungulate body part profiles will be addressed in Chapter 6.
Observable mammalian carnivore damage includes crenellation, salivary rounding of broken
edges, punctures, tooth drag marks, evidence of digestion, and an absence of long bone
epiphyses in conjunction with other damage (Binford 1981:44-49, 51-77; Brain 1981;
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Marean and Spencer 1991; Stiner 1994:140-147). Presence or absence and type of carnivore
gnawing was recorded during the analysis of the Klissoura Cave 1 faunal assemblages.
Rodents can be involved in both the accumulation and destruction of bone. In
particular, wood rats (Neotoma sp.) are common accumulators of small mammal, fish and
reptile bones, as well as medium-sized ungulates (Hockett 1989; Hoffman and Hays 1987).
Porcupines (Hystrix africaeaustralis, the African porcupine, in particular) are also collecting
agents, and they often cause major damage to ungulate remains by gnawing them (Brain
1980, 1981; Maguire, et al. 1980). Though these particular species were not necessarily
present at Klissoura Cave 1 and therefore did not disrupt the assemblages, other rodents were
in the area and their impact on the faunas must be evaluated. To do this, rodent damage was
recorded when present on a specimen.
Diurnal raptors (order Falconiformes) and owls (order Strigiformes) are potential
accumulators of bird, mammal, and reptile remains in an archaeological assemblage. Many
raptors and owls use rock faces and shelters for nesting (Hockett 1996; Lloveras, et al.
2008a). A series of taphonomic studies were undertaken to examine the effects of Falco
rusticolus (gyrfalcon), Aquila heliacal (imperial eagles), Asio otus (long-eared owl), Strix
aluco (tawny owl), and Bubo bubo (eagle owl) feeding patterns and digestion of smaller
avian prey (Bocheński, et al. 1997; Bocheński and Tomek 1994; Bocheński, et al. 1993).
These studies indicate that fragmentation rates and the survivorship of particular elements
differ depending on the prey species. Taphonomic signatures of specific avian predators may
be difficult to discern in an archaeological assemblage, which may represent a palimpsest of
many raptor meals. However, damage from digestion is a common feature of the
assemblages that many predatory birds accumulate: snowy owls 6-60% of NISP, gyrfalcons
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67-97%, long-eared owls 55%, tawny owls 82% and eagle owls 84% (damage percentages
for the last three raptors only includes femurs) (Bocheński, et al. 1997; Bocheński and
Tomek 1994; Bocheński, et al. 1993). If Falconiformes or Strigiformes were the
accumulators of bird remains, some digestion damage should be apparent. Similarly, leporid
remains should also be expected to exhibit corrosion if they were digested and regurgitated
by diurnal raptors or owls (Hockett 1996; Lloveras, et al. 2008a, 2008c). Observations of
Falconiformes and Strigiformes nesting sites in North America, Europe and Africa indicate
leporid assemblages accumulated by eagles tend to contain mostly hind limbs, and
occasionally head parts (Cruz-Uribe and Klein 1998; Hockett and Haws 2002; Lloveras, et
al. 2008c). Owl pellets are identified as containing more forelimbs and head parts (Hockett
and Haws 2002), but owl nesting sites are observed to have the opposite set of remains
(Lloveras, et al. 2008a). Therefore, hare body part profiles dominated by either front limbs or
hind limbs may be indicative of raptor nesting. On the other hand, a combination of diurnal
raptor and owl feeding behaviors may create the illusion of a more even representation of
forelimb and hind limbs in an assemblage, so again digestion damage is necessary for
implicating birds of prey in contributing to the accumulation of faunas in an archaeological
site. Sampson (2000) analyzed tortoise bones from multiple South African rock shelters and
raptor kill sites in order to distinguish between the taphonomic signatures of diurnal raptors,
small carnivores and humans. Raptor roosting sites contain higher proportions of intact
tortoise crania and axial elements, low frequencies of forelimbs, and few carapace or plastron
fragments, since shells tend to be abandoned at kill sites. Small carnivores assemblages, on
the other hand, tend to contain few cranial and axial elements, and larger numbers of
forelimbs and shoulder girdles (Sampson 2000). The major differences between human and
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non-human assemblages are a high degree of burning and higher percentages of shell
fragments in anthropogenic contexts (Sampson 2000).
Another set of taphonomic processes that may alter an archaeological assemblage is
weathering, defined by Behrensmeyer (1978:153) as the process of separation and
destruction of the original organic and inorganic components of bone by physical and
chemical agents. Weathering encompasses a range of characteristics, from cracking and
exfoliation, to root damage, to chemical diagenesis. One set of weathering processes, that
includes bleaching and erosion, occurs more frequently at open-air sites when remains are
exposed to the sun and wind before being buried (e.g., Behrensmeyer 1978; Lyman and Fox
1989). In order to identify surface weathering at Klissoura Cave 1, several different
weathering stages for each of the specimens are recorded (following Stiner 2005, but also
including root damage).
Soil chemistry is also a major factor in the preservation of bone. A number of studies
focusing on Paleolithic cave sites have identified the presence of specific minerals in
sediments, specifically calcite and dahllite, as indicators of good bone and wood ash
preservation (Karkanas 2001; Karkanas, et al. 1999; Schiegl, et al. 1996; Stiner, et al. 2001;
Weiner, et al. 1993). Conversely, an absence of calcite and dahllite indicates that bone should
not have preserved archaeologically. This principle was applied to Theopetra Cave in
northern Greece, and Hayonim and Kebara Caves in Israel to explain the lack of faunal
materials in certain areas of the sites (Karkanas, et al. 1999; Schiegl, et al. 1996; Stiner, et al.
2001; Weiner, et al. 1993). To the extent that distributions of faunal remains (see Chapter 7)
are heterogeneous at Klissoura Cave 1, particularly as compared to the frequency of lithics,
in situ dissolution should be investigated.
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A final taphonomic process that will be addressed here is density-mediated attrition,
which is bone loss as a function of the structural density of bone macrostructures. The
structural density of an element portion is the ratio of the mass of the bone to its volume
(Lyman 1984:264; 1994:237). The basic premise of density-mediated attrition is that portions
of elements with low structural density, for example highly porous long bone epiphyses,
should be more susceptible to being damaged or destroyed by mechanical and chemical
processes than are portions of elements with high structural density, such as long bone shafts
(Binford and Bertram 1977; Brain 1981; Lyman 1994). Mechanical (and possibly chemical)
attrition should disproportionately affect specimens with low bulk densities because they
have high surface area to volume ratios and more exposed surface to be damaged; therefore,
specimens with low bulk density should have reduced survivorship in an archaeological
assemblage (Lyman 1984:279; 1994:239). Various methods including photon densitometry,
duel energy X-ray densitometry, and computed tomography (CT) have been used to derive
bone density standards for comparison to archaeological data (Kreutzer 1992; Lam, et al.
1999; Lyman 1984, 1994). Bone density values have been derived for diverse taxa, including
white-tailed deer, caribou, pronghorn, domestic sheep, cattle, pigs, leporids, marmots, deer,
and bison, among others (Ioannidou 2003; Kreutzer 1992; Lyman 1984, 1992; Pavao and
Stahl 1999; Symmons 2005). Though this list is extensive, it does not include all of the
important species in Mediterranean archaeological assemblages. Some criticisms have been
leveled concerning the use of bone density values of one species as a proxy for other species
in an archaeological assemblage because of recorded variation between taxa that stems
largely from locomotive adaptations or the way that mass is distributed across the bodies of
different taxa (Ioannidou 2003; Kreutzer 1992; Pavao and Stahl 1999). However, bone
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density values of similar species are routinely applied to archaeological assemblages, because
tests of density-mediated attrition are so valuable and in some cases there are no other sets of
data to use (Stiner 2005:86). Some degree of ambiguity in the reference data used in this
study cannot be avoided. However, Lam et al. (1999), among others, have argued that
differences are mainly between mammalian families rather than within them, or are the result
of studies using different methods to derive density values. They compare bone density
values for bovids, cervids, and equids and find that intertaxonomic variation is low enough
that density values can appropriately be applied to different species with similar
morphologies (Lam et al. 1999:356). Therefore, reference data derived for other cervids (i.e.,
Odocoileus sp.) are generally suitable for analyzing fallow deer survivorship at Klissoura
Cave 1.
Much of the work seeking to understand differences in the structural density of bone
and density-mediated attrition was stimulated by Binford’s (1978) utility curve study in his
ethnoarchaeological work with the Nunamiut (see Chapter 6). Utility curves have
subsequently been widely applied to faunal assemblages in order to understand transport
decisions by past foragers. However, skeletal portions that rank high in utility because of
their meat or fat content tend to rank low in structural density (Lyman 1985), so in cases
where an assemblage suffered from density-mediated attrition, the representation of remains
fit with the interpretation of people transporting carcass portions based on reverse utility
strategies (e.g., selectively transporting low-utility elements).
Bone loss from density-meditated attrition can also arise from chemical and
mechanical weathering processes, as well as carnivore gnawing and human butchery (e.g.,
Binford and Bertram 1977; Davis 1987; Lyman 1984). The most porous, and therefore least
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dense, portions of elements are filled with deposits of bone grease. This grease is attractive to
carnivores, which is why they tend to preferentially ravage long bone epiphyses and other
porous elements (Blumenschine 1988; Marean and Spencer 1991; Marean, et al. 1992), and
is also exploited by some humans during bone grease rendering (e.g., Binford 1978; Brink
1997; Lupo and Schmitt 1997; Vehik 1977). Therefore, evidence of density-mediated
attrition only indicates that bone loss related to structural density occurred in an
archaeological assemblage, but it does not identify the cause as being non-human chemical or
mechanical weathering, carnivore gnawing, or human butchery or transport patterns. An
evaluation of density-mediated attrition nonetheless is crucial to interpretations concerning
transport decisions.
Two tests evaluate the potential of density-mediated attrition to explain prey body
part representation at Klissoura Cave 1. The first compares the highest tooth-based MNE to
bone-based MNE for skull parts in ungulate species in each archaeological layer (following
Stiner 1994:99-103) and is based on comparable differences between bone and tooth
accumulation and attrition. In the transport of ungulate remains to an archaeological site, it is
expected that teeth would tend to remain with the skull, which would lead to a roughly 1:1
ratio of tooth and cranial bone-based MNE. Overall, mammal teeth are less susceptible than
bone to most destructive processes because the mineral component of tooth enamel (95%) far
exceeds that of any bone (70%) (Currey 1984; Hillson 2005:146; Lyman 1984:72, 79). If
significant fragmentation occurred during the processing of the heads by human hunters, or if
gnawing or other attritional processes happened post-depositionally, even durable bony
features on the skull should break down more often than teeth. Therefore, a significant bias in
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the ratio of tooth and cranial bone-based MNE in favor of higher tooth-based MNE would
indicate some attritional processes affected the assemblages.
The second test of density-mediated attrition compares observed patterns in bone
representation in an assemblage to independent standards of bone tissue density values
obtained by the photon-densitometry technique (Lyman 1984; 1994: see also Lam et al. 1999
on the CT technique). Bone survivorship is evaluated for each portion of an element by
dividing the observed frequency of each portion in an assemblage by the expected frequency
for this part in a complete skeleton. This value is then standardized against the most
commonly represented body part in the assemblage for a given species to determine percent
survivorship (Binford 1978; Lyman 1994). Percent bone survivorship was determined for
fallow deer and hare remains because they are the most common taxa through the sequence.
In this test, and in the discussion of body part representation in Chapter 6, hare data are
combined with data for indeterminate small mammals since most of these remains probably
come from hare at Klissoura Cave 1, as opposed to similarly-sized animals such as fox or
wild cat which are extremely rare in the assemblages. Similarly, remains designated as
medium ungulates were combined with taxon-specific fallow deer data, since fallow deer
was always the most common medium ungulate species in the faunas by a large margin. In
the Aurignacian layer IV assemblage, where there was a significant proportion of other
medium ungulates (i.e., ibex), not all remains that could only be identified as “medium
ungulate” were lumped with fallow deer data as described previously. Rather, the proportion
of medium ungulates specifically identified as fallow deer and ibex in this layer was noted
(57.4% and 42.6%, respectively, based on NISP) and 57.4% of elements identified as
“medium ungulate” were lumped with the fallow deer data. This is hardly ideal, particularly
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because transport decisions for ibex and fallow deer may have been different if ibex were
hunted further from the site, but it was done in order to avoid over-representing fallow deer
remains. Bone density standards for American deer (Odocoileus sp.) were applied to fallow
deer from Klissoura Cave 1 (from Lyman 1982, 1984), and the standards applied to European
hare were those developed for snowshoe hare (Lepus canadensis) (Pavao and Stahl 1999).
Comparing observed and expected body part representations at an archaeological site
is useful for identifying biases in zooarchaeological assemblages, but it cannot determine
whether human or non-human processes led to the biases. The general type of site must be
identified (e.g., hunting station, processing site, residential camp) as well as the agent of
transport and modification. Ethnographically, human foragers are shown to be selective
about which ungulate body parts will be carried over long distances, whereas small game
carcasses tend to be carried to a site whole (e.g., Binford 1978; Bunn, et al. 1988; O'Connell,
et al. 1988; Schmitt and Lupo 1995; Yellen 1991). Once human foragers accumulate these
remains at a site, the prey items would be subject to processing for consumption as well as
post-depositional processes. Transport biases aside, the survivorship of hare and fallow deer
bones should be about the same at Klissoura Cave 1 if post-depositional destruction was the
main cause of anatomical bias (e.g., Manne and Bicho 2009; Manne, et al. 2005; Munro
2004). If hare bones show no indications of density-mediated attrition but the ungulate
remains do, then the case can be made that biases in fallow deer body part representation
were caused by selective transport or intensive processing on-site (following Munro 2004).
Conversely, several explanations are possible if there is evidence that hare bones underwent
density-mediated attrition but ungulate bones did not. One is that in general, hare elements
are structurally less dense than deer bones (Pavao and Stahl 1999; see values in Lyman 1994)
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and are more susceptible to trampling damage. Another possible explanation is that hares
were not brought to the site whole, or they were processed and consumed in different areas of
the site (e.g., Cochard and Brugal 2004). These hypotheses can be tested to some extent by
comparing survivorship rates of small mammals and medium ungulates in a given
assemblage, and will be explored further in Chapter 6.
In discussing survivorship of different bone elements, the issue of shaft versus endbased MNE is also addressed. It is assumed in this analysis that in most cases, when large
animals were butchered in the field, carcasses were partitioned in such a way that most bone
shafts were accompanied by at least one epiphyses when transported to the home base.
Therefore, it is expected that accumulated remains, barring intense butchery or postdepositional attrition, should include equivalent ratios of proximal and/or distal long bone
ends, and shafts, for each appendicular element. In certain cases, examining the ratio of long
bone ends to shafts may indicate attritional processes of noncultural or cultural types. In
general, based on structural density alone, a slight bias (1:2 or 1:3) of cancellous to compact
elements may be expected (using values from Lyman 1984; Lyman 1994; see also values
from Lam et al. 1999 in Stiner 2004). A more extreme ratio of 1:8 cancellous to compact
element survivorship was estimated from bone ravaging experiments with hyenas (Capaldo
1997; Marean and Spencer 1991; Marean, et al. 1992), refitting studies of Paleolithic
assemblages (Marean and Kim 1998), and measurements of limb shaft densities using
adjusted computed tomography (Lam, et al. 1998; Lam, et al. 1999). Ratios of ends (based on
the maximum from the proximal or distal end) to shafts were recorded for the different layers
of Klissoura Cave 1. As with the other tests discussed here, only medium ungulates were
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taken into account because they have the largest sample size through the sequence. The limb
elements considered were the humerus, radius, ulna, metacarpal, femur, tibia and metatarsal.
The suite of analyses discussed above can, taken together, provide insight into
understanding and interpreting the condition of the faunal remains at Klissoura Cave 1. Once
non-human taphonomic biases are taken into account, hominin prey selection, transport
decisions, and butchery patterns can be more fully understood.
RESULTS
Input Analysis and Fragmentation
An analysis of faunal input (or density) of Klissoura Cave 1 indicates that there is not
a relationship between sample size and faunal input, once excavated volume is taken into
account (Table 4.2, n = 14, rp = 0.326, p = 0.255). The layers with the smallest sample sizes
(Mesolithic 3-5a, Epigravettian IIa-d, Early Upper Paleolithic V, and Middle Paleolithic X)
also have some of the lowest densities of animal remains (<0.50 NISP/L), but larger samples
from the Middle Paleolithic layers XI-XIV and XV-XVII also have extremely low rates of
faunal input. Middle and Upper Paleolithic layers with comparatively moderate input rates
(0.51-.90) include Upper Paleolithic layer III’, Middle Paleolithic layer XXa-XXb.
Aurignacian layers (IV, IIIe-g, IIIb-d) and the non-Aurignacian Upper Paleolithic (III”) have
the highest rates of faunal input, and there is a trend toward increasing density through time.
Middle Paleolithic layers VIII and XVIII-XIX also have high faunal densities. In general,
this follows the geoarchaeological evidence quite well (Karkanas 2010), with the largest
anthropogenic signature from a faunal perspective occurring mainly in the earlier Upper
Paleolithic layers, though use of the site or preservation clearly declines in later periods
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(Table 4.2). Not surprisingly there is no correlation between input rate and time period (n =
14, rs = 0.033, p = 0.911) which suggests that there was not a steady increase in the use of
Klissoura Cave 1 over the years; rather the intensity or duration of occupation reached an
apex during the Aurignacian.
Mean length and standard deviations for different body classes in each archaeological
layer are reported in Table 4.3. Unsurprisingly, the average fragment size for each body class
seems to increase as body size increases, but differences in sample sizes make comparisons
between body class difficult to quantify. There is no temporal trend of fragmentation size for
the overall sample (n = 14, rs = -0.103, p = 0.725) or medium ungulates, the most common
prey type at the site (n = 14, rs = -0.402, p = 0.154); no change in fragmentation through the
assemblages is apparent. Initial fragmentation was probably related to carcass butchery, but
subsequent fragmentation can likely be attributed to trampling or shattering caused by high
temperatures as hearths were built on bone-rich surfaces (see Chapter 6). The MNE/NISP
fragmentation index is presented in Table 4.4. As with mean length, there is no trend between
time and fragmentation index value (n = 13, rs = -0.249, p = 0.411). The index value from the
Epigravettian layer (IIb-d) is excluded from the correlation because the sample is small. In
addition to the lack of temporal trend for either indication of fragmentation, there is also no
correlation between the length of medium ungulate specimens and fragmentation index (n =
13, rp = -0.039, p = 0.899). This means that layers with higher rates of fragmentation do not
necessarily have the smallest specimens.
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Table 4.3 Mean fragment length for different body classes, by cultural layer.
Carnivore Damage and Weathering
Evidence of carnivore activity is extremely rare in the Klissoura Cave 1 assemblages.
A range of small and large carnivore species were identified in the faunas (Chapter 5), but it
seems they did not use the site for shelter. No unfused or neonate carnivore remains were
found, suggesting that the site was not used as a den. Three hyena coprolites, one in the nonAurignacian Upper Paleolithic (III”), the others in Middle Paleolithic layers (VIII and X)
indicates that hyenas visited the cave, but this was an extremely rare occurrence. Only thirty-
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Table 4.4 MNE:NISP index values for medium ungulate cranial elements and all elements in the
Klissoura Cave 1 assemblage. *Note small sample sizes.
five specimens from the entire assemblage have evidence of carnivore tooth puncture or drag
marks, and these frequencies are far too low to elucidate any kind of trend (Table 4.5).
Finally, ungulate body part profiles (Chapter 6) are similar to transport patterns of humans,
not large carnivores (Stiner 1991, 1994). Incidences of rodent damage are even rarer than
carnivore damage in the Klissoura Cave 1 assemblages, with just three specimens from all
layers displaying rodent gnawing (Table 4.5).
There is little evidence indicating that diurnal raptors or owls were major
accumulative agents in the assemblages. Though several raptor species, specifically Aquila
chrysaetos (Golden eagle), Tyto alba (barn owl), Bubo bubo (Eurasian eagle owl) and Asio
sp. (eared owls) were identified in the Klissoura Cave 1 assemblages (Bocheński and Tomek
2010), their frequencies are exceptionally low (NISP<8 for the entire Upper Paleolithic and
later periods). No cranial puncture marks or digestion damage was apparent on any of the
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Table 4.5 Frequencies of gnawing and weathering damage on faunal specimens by cultural layer.
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small game remains. It is possible that some of the micromammals were the remains of owl
or raptor meals, but these small creatures were excluded from this study to be analyzed by
specialists for environmental reconstructions. Additionally, hare body part analysis does not
include a high frequency of hind or front limb bones (Chapter 6), which may be expected in
an assemblage accumulated by avian predators. Tortoise remains have high incidences of
burning and are mostly carapace and plastron fragments, consistent with human exploitation.
Weathering was not a major taphonomic process at Klissoura Cave 1. Cracks and
exfoliation that often accompany exposure to the sun are almost completely absent from the
assemblages (Table 4.5). Chemical weathering was the most frequently observed process,
which often made it appear that the specimens had been burned or etched by acid (Figure
4.3). This type of weathering was the most common in the Middle Paleolithic layers and is
almost absent from the Upper Paleolithic and later layers, though overall it is still fairly rare
(< 2.5% in any layer) (Table 4.5).
Figure 4.3 Chemical weathering on Middle Paleolithic long bone fragment. From layer XIV.
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Density-mediated attrition
Ratios of ungulate tooth to bone-based cranial MNE are presented in Table 4.6
(Figure 4.4) by layer. All ungulate taxa were combined to create a more robust sample
(Appendix C contains expanded data). Of the fourteen archaeological layers, four of the
samples are too small (MNE<6) to draw conclusions related to density-mediated bias. Of the
remaining layers, most have near-even ratios of tooth to cranial bone-based MNE, with a few
exceptions. Aurignacian layer IV and Middle Paleolithic layers VIII have tooth-based MNE
counts that are double the cranial bone-based MNE (Table 4.6). Middle Paleolithic layer VIII
seems to have suffered from some density-mediated attrition based on bulk bone density tests
(see below), but there is no explanation for Aurignacian layer IV. One possibility is that
crania were more smashed up in this layer. This should be testable, however, by comparing
the fragmentation index for cranial parts (Table 4.4) with the proportion of tooth to cranial
bone-based MNE (Table 4.6). A higher tooth to bone-based MNE ratio reflects poor
survivorship of cranial bones, and a low index value for cranial fragmentation (MNE/NISP)
indicates a higher degree of breakage. Therefore, it might be expected that tooth to cranial
bone-based MNE ratios should correlate negatively with the fragmentation index if increased
processing or post-depositional fragmentation were to blame for an under-representation of
cranial bones in the assemblages. This is not, however, the case at Klissoura Cave 1; a
Pearson’s correlation indicates no relationship between the cranial fragmentation index and
the proportion of tooth to cranial bone-based MNE (n = 9, rp = -0.283, p = 0.461). It seems
unlikely, therefore, that high the proportion of tooth to bone-based MNE in Aurignacian
layer IV is due to fragmentation of cranial bones, and this anomaly cannot be readily
explained. The non-Aurignacian Upper Paleolithic layer III” and Middle Paleolithic layers
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XVIII-XIX have much higher bone than tooth-based MNE counts. This may relate to greater
tooth fragmentation rates in these layers based on excavation damage or post-depositional
processes, but this is unclear. Neither of these layers have abnormally high fragmentation
rates overall (Table 4.4).
Table 4.6 Tooth and bone-based MNE for all ungulates by layer.
Figure 4.4 Proportion of tooth to bone-based MNE for all layers at Klissoura Cave 1. Data from
Table 4.6.
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Turning to bulk density, the results of a Spearman’s rank-order correlation between
bone density values and percent survivorship for hares and fallow deer/medium ungulates
from each layer at Klissoura Cave 1 are presented in Table 4.7. Note that the Epigravettian
(IIb-d), Early Upper Paleolithic (Uluzzian layer V) and Middle Paleolithic (X) are excluded
from the comparison because of small samples for both hare and fallow deer. Likewise,
fallow deer values from the Mesolithic (3-5a), and hare values from the non-Aurignacian
Upper Paleolithic industry (III”) and the entire Middle Paleolithic are unavailable due to the
low frequency of hares in these assemblages. The hare remains from the Mesolithic (3-5a)
and Mediterranean backed-bladelet industry (III’) display a significant, positive relationship
between bone density and percent survivorship. The rs2 values, however, indicate that
density-mediated attrition has the potential to explain only eight to sixteen percent of the
variation in skeletal survivorship from these levels (Table 4.7). For fallow deer in the Upper
Paleolithic and later layers, only the middle Aurignacian (IIIe-g) indicates a significant
positive relationship between percent survivorship and bone density, and here densitymediated attrition can explain no more than five percent of the variation in deer body part
representation (Table 4.7). In addition, hare survivorship does not display a correlation with
bone density for this layer, even though hare remains are on a whole more fragile than
ungulate bones. In the Middle Paleolithic there is a positive correlation between percent
survivorship and bone density values for fallow deer in all of the layers, and rs2 values
indicate that density-mediated attrition has the potential to explain 6-26% of the variation in
skeletal survivorship in these layers (Table 4.7).
Maximum MNE based on long bone ends and shafts for selected long bones from
medium ungulates in the Klissoura Cave 1 assemblages are presented in Appendix D.
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Table 4.7 Spearman’s rank-order correlation between percent survivorship and bone density values
for European hare and fallow deer/medium ungulate remains. Data from Appendices A and B.
*Significant values
Values for different elements are collapsed into all long bone ends and shafts in Table 4.8
and are plotted in Figure 4.5. A 1:1 ratio is expected when shafts and ends are equally wellpreserved because values are used for the end with the highest MNE. In almost all layers,
except for the Middle Paleolithic (XVIII-XIX), MNE values for long bone shafts and ends
are either equal, or end-based MNE is much higher than shaft-based MNE. A closer look at
the data reveals that most of the layers with higher end-based MNE values are from the
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Upper Paleolithic layers, while the Middle Paleolithic layers have nearly a 1:1 ratio between
shaft and end based MNE (Table 4.8). Potential explanations for this are discussed below.
Table 4.8 End and shaft-based MNE values for medium ungulate long bones by cultural layer.
Elements include humerus, radius, ulna, metacarpal, femur, tibia and metatarsal.
Figure 4.5 Proportion of shaft-based and end-based MNE for all layers at Klissoura Cave 1. Data
from Table 4.8.
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DISCUSSION AND CONCLUSIONS
Several interesting points concerning site use and the condition of faunal remains at
Klissoura Cave 1 are apparent in the results presented above. Perhaps the most striking
feature is the extent to which the faunas were primarily accumulated and modified by
Pleistocene hominins. This bodes well for interpretations of small game use and human
transport and butchery patterns presented in later chapters. Fragmentation rates of faunal
materials indicate no temporal trend, so they cannot be taken to indicate an increase or
decrease in the intensity of processing activities or other sources of mechanical damage over
time. The high degree of fragmentation at Klissoura Cave 1 is probably largely due to postdepositional factors, such as trampling or compacting by sediments, though relationships
between fragment size and butchery intensity will be explored in Chapter 6.
In terms of site occupation intensity, rates of faunal input support geoarchaeological
conclusions set forth by Karkanas (2010), who interprets the Middle Paleolithic,
Epipaleolithic and Mesolithic sediments as having a higher natural input and the Upper
Paleolithic layers as having a more intense anthropogenic signature. Occupation intensity can
refer to the duration of time a site was used, how often it was used, or how many people were
utilizing the site at a given time. Distinguishing between these patterns of use in the
archaeological record is difficult, particularly because an increase in occupation intensity
could represent something as minor as ten people as opposed to five using a site for a week,
or a site being occupied for weeks rather than days. In this context, it simply means a general
increase in site use, which could encompass any of these behaviors. According to faunal
density data, the site was occupied most intensively during the Aurignacian and the overlying
non-Aurignacian Upper Paleolithic (III”). The Aurignacian layers (IV, IIIe-g) of Klissoura
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Cave 1 also contain dozens of clay-lined hearth features, and a possible rock-delineated
structure (Karkanas, et al. 2004; Koumouzelis, Ginter, et al. 2001; Pawlikowski, et al. 2000;
Stiner 2010). There was a subsequent decline in occupation intensity, particularly following
the depositional hiatus of the Last Glacial Maximum (Karkanas 2010). This decline may
indicate the use of different kinds of sites or different uses of the landscape in later periods,
perhaps even other caves in Klissoura Gorge.
The scarcity of carnivore remains, evidence for carnivore denning, and modification
of the faunas is striking, as it is clear that carnivores were present and active on the late
Pleistocene landscape in Southern Greece (Gamble 1986; Payne 1975; Reisch 1976; Stiner
and Munro 2011; Tsoukala 1992). It is possible that the Klissoura Gorge with its many caves
provided ample options for sheltering carnivores, so that the occasional human presence at
Klissoura Cave 1 was enough to discourage carnivores from using the site. Another option is
that carnivores used the deeper, narrower parts of the cave for shelter, which is an area that
was not included in the excavation. If this were true, we might still expect to see appreciable
frequencies of carnivore damage on the archaeological assemblages, which is certainly not
the case. Evidence for modifications made by birds of prey are also rare in the faunas, as is
rodent gnawing. Surface weathering of faunal remains was not a factor in the Klissoura Cave
1 assemblages. Also, there is no indication of differential bone loss in specific parts of the
site (see Chapter 7) so chemical diagenesis was probably not a major problem either.
Results of tests evaluating density-mediated attrition are somewhat mixed, though
evidence of density-mediated bias does not indicate the underlying cause (i.e. human or nonhuman) of attrition. Overall, deer bone survivorship is uncorrelated or only weakly correlated
with bone density in the Upper Paleolithic layers. The cranial bone-based MNE nonetheless
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is much lower than expected as compared to tooth-based MNE for Aurignacian layer IV; this
discrepancy cannot be explained by in situ attrition or heavy fragmentation of cranial parts.
Hare bone survivorship does correlate with bone density in the Mediterranean backedbladelet industry (III’) and above, so there may be some density-mediated processes at play
in these later layers. A lack of correlation between deer/medium ungulate bone survivorship
and bone density in layer III’, however, suggests that any density-mediated processes in the
upper layers may have only affected the structurally least dense or fragile small elements, in
this case hare remains. A discussion of body part representation in Chapter 6 will allow us to
evaluate some of the potential variation in bone survivorship as a product of human behavior.
Correlations between percent survivorship and bone density for ungulate remains in
the Middle Paleolithic elicit several questions concerning game use in these earlier periods.
1) Are fallow deer elements missing in the Middle Paleolithic layers because of densitymediated attrition or transport decisions by human foragers? 2) Are hare remains lacking in
this lower layers of Klissoura Cave 1 because they were not exploited by Middle Paleolithic
hunters, or are they absent due to attritional processes? First, it must be pointed out that there
is only a bias in tooth to cranial bone-based MNE in the latest Middle Paleolithic layer (VIII)
(Table 4.6). At the same time, there is actually a greater representation of cranial bone-based
MNE compared to tooth-based MNE in Middle Paleolithic layer XVIII-XIX. It seems likely,
then, that though there is some correlation between body part representation and skeletal
density in the Middle Paleolithic, some of the differential representation of anatomical parts
was influenced by human transport decisions or butchery patterns, which can also relate to
density-mediated processes (see Chapter 6). As for hare remains in the Middle Paleolithic
layers, if density-mediated attrition was a factor, we would expect to find only the most
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dense elements. Despite the fact that sample size is small, hare remains that are present in the
Middle Paleolithic layers span an entire range of structural densities, including many that are
far less dense that any fallow deer elements (Appendix E).
An interesting and rather counterintuitive bias in the MNE of under-represented long
bone shafts versus over-represented ends was observed in several of the assemblages,
particularly in the Upper Paleolithic layers. The most likely explanation for this is the
macroscopic condition of the Upper Paleolithic remains. Most of the Upper Paleolithic
faunas were covered with an ashy layer that obscured many macroscopic markings on the
specimens. It is likely that this same ash also hid diagnostic shaft features, such as nutrient
foramina and muscle attachments. This provides yet another argument in favor of relying on
both long bone shaft and end features for analyzing zooarchaeological assemblages, which is
already done by most faunal analysts (Bar-Oz 2005; Binford 1978; Brain 1981; Bunn and
Kroll 1986; Delpech 1998; Morlan 1994; Rogers 2000; Stiner 1991, 1994, 2005; Todd and
Rapson 1988), because it is never clear at the outset what kind of analytical biases may be
encountered in an assemblage.
Though some non-human taphonomic processes affected the faunas of Klissoura
Cave 1, they were very minor overall. Based on the discussion outlined in this chapter it
seems unlikely that broad patterns of human behavior related to changes in species use,
transport decisions or butchery patterns were obscured by non-human taphonomic processes
acting on the assemblages.
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CHAPTER 5: RELATIVE SPECIES ABUNDANCE
AND DIVERSITY OF PREY TYPES
INTRODUCTION
Changes in human subsistence strategies as reflected by prey choice over a long
archaeological sequence are informative of changing environmental conditions, shifts in
human demography, or as we see at Klissoura Cave 1, both. Using multiple methods,
including analyzing prey groups based on body size and flight behavior (following Stiner, et
al. 2000), tests of species diversity, ratios of prey types, biomass comparisons, and input
analysis (see Chapter 4), this chapter evaluates diachronic change in prey choice from the
Middle Paleolithic through Mesolithic at Klissoura Cave 1. Following predictions laid out in
Chapter 3, it is expected that some of the changes in the species representation of large game
animals, as well as great bustard, track environmental changes. Conversely, shifts in the
overall composition of the small game component, and changes in the proportion of large and
small game through time, relates more closely to increased occupation intensity at the site or
in the region as a whole.
METHODS
Several different approaches are employed to evaluate prey choice over time at
Klissoura Cave 1. Number of identified specimen (NISP) counts for all species identified in
the eight Upper Paleolithic and later layers and six Middle Paleolithic units are presented
(Appendix F). The taxa are divided into eight broad prey categories based on body size and
behavioral characteristics; there are four different size classes for ungulates, small and large
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carnivores, small slow-moving game, and small fast-moving game (Table 5.1). It is predicted
that human foragers would rank animals according to differences in their handling costs and
return rates, with large-bodied ungulates being preferable (see Chapter 1, Table 5.1).
Carnivores are considered separately because they are behaviorally different than other small
and large game, and are very uncommon at Klissoura Cave 1 though they represent human
prey (as opposed to dying naturally at the site). The small, slow-moving category includes
tortoises (Testudo sp.), which are easy to collect by foragers of all ages and conditions. The
small, fast-moving game category includes hares (Lepus europaeus) and birds of various
sizes (most notably great bustard, Otis tarda, and rock partridge, Alectoris graeca), species
that tend to require a significant technological investment (e.g. snares or nets) in order to be
exploited efficiently (Cannon 2000; Jones 2006; Madsen and Schmitt 1998, but see also
Ugan 2005 for differences between terrestrial game, fish and invertebrates). Class
designations including small birds (<0.1 kg) and indeterminate snakes were excluded from
the groupings because it is not certain that they were exploited by humans; however, they
exist in such low frequencies (NISP<15 in the entire assemblage) that their inclusion would
have little effect on these interpretations.
The four major small game species found at Klissoura Cave 1 have very different life
history characteristics, habitat requirements, and flight responses; the bustard is further
distinguished by its large size (see Table 5.1). Tortoises have low capture costs and high
return rates because they are slow-moving; therefore, they are expected to be preferred prey
when available (Stiner, et al. 2000). They are a slow-growing species, taking a long time to
reach reproductive maturity and continuing to grow throughout their lifetimes. They are
sexually dimorphic in terms of size, with females tending to be larger (Blasco, et al. 1986-87;
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Hailey, et al. 1988; Lambert 1982; Swingland and Stubbs 1985). Therefore, the most obvious
individuals on the landscape, and the most desirable for human collection, are the large, older
females that are the reproductive base of the population (Stiner, et al. 2000). Climatic
variation would never have been great enough to eliminate Mediterranean tortoises from
southern Greece; they are found in human diets throughout much of the Middle and Late
Table 5.1 Taxa included in each body class category and their respective weight ranges and predicted
ranking. Mass values from Nowack (1999) and Silva and Downing (1995).
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Pleistocene in the eastern Mediterranean and Middle East (Reynaud Savioz and Morel 2005;
Speth and Tchernov 2002; Stiner 2005, 2009; Stiner, et al. 2000). Therefore, a decrease in
tortoises in an archaeological sequence may be the result of overexploitation by human
foragers.
Though they are categorized as small, fast-moving animals, great bustards potentially
have higher return rates than either hares or partridges because they are appreciably larger
(Table 5.1). Like tortoises, however, bustards have low rates of reproduction and
development and rely disproportionately on older females to perpetuate the species (Morales,
et al. 2002). Females over six years of age are twice as successful at reproducing as younger
birds, even then they average only 0.40 chicks per year and are unlikely to breed two years in
a row (Morales, et al. 2002). Successful individuals tend to be long-lived and population
turnover rates are comparatively low; in this sense bustard populations are similar to tortoises
in that they are easily stressed with the removal of a few older females. Heavy exploitation of
either of these taxa could not be sustained for long.
Rock partridge and European hare likely have similar return rates to one another,
which are lower than the other two small game taxa, because they have rapid flight responses
when confronted by a predator. Both species are prolific and are able to reproduce in the first
year of life. Partridges have yearly clutch sizes of between twelve and eighteen eggs
(Vavalekas, et al. 1993), and hares give birth to one to four litters per year of up to five
leverets (Burton 1991). Their productivity as modeled against Mediterranean tortoises can be
as much as seven times greater (Stiner, et al. 2000).
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Examining richness and evenness (i.e. diversity) in a faunal assemblage is a common
way to evaluate relative species abundance and to control for sample size differences in
zooarchaeological assemblages (Grayson and Cannon 1999; Grayson and Delpech 1998;
Nagaoka 2001). There are many tools for analyzing species diversity, and various terms have
been used in different ways in ecological and anthropological literature (see Lyman 2008:175
for a review). In this presentation, taxonomic richness refers to the number of taxa identified
to the genus level, such that an assemblage with ten different taxa is richer than an
assemblage with eight provided that sample sizes are similar. Genus is used rather than
species because the species is not altogether certain for some taxa such as Capra (though it is
most probably C. ibex). Additionally, there may have been up to three species of similarsized tortoises (Testudo graeca, T. hermanni, and T. marginata) in southern Greece and the
remains could only be identified to Testudo sp. In the case of diversity indexes, the level at
which taxonomic identifications are made is less important than maintaining consistency
with the taxonomic level at which they are tallied (Lyman 2008:174). A related method to
evaluate species abundance is taxonomic evenness, which measures how individuals
distribute across different categories, or taxa (Smith and Wilson 1996). Taxonomically even
assemblages have a similar number of individuals for each of the different genera. One
technique for evaluating the evenness of archaeofaunas is using the reciprocal of Simpson’s
Index (Grayson and Delpech 2002; James 1990; Jones 2004; Schmitt and Lupo 1995; Stiner
2001). It is calculated as 1/D, or 1/Σ(ρi)2, where ρ is the proportion of taxon i in the total
assemblage (Levins 1968; Simpson 1949). A lower value for 1/D indicates the
disproportionate representation of one taxon. This simple index evaluates species richness
and evenness simultaneously and corrects for sample size differences among layers in an
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archaeological assemblage. Prey diversity, as it relates to the input analysis introduced in
Chapter 4, is also examined.
Presenting the minimum number of individuals (MNI) is a common technique in
faunal analyses (Grayson 1984). Like many useful zooarchaeological counting units, it is
derived from NISP, though it suffers more than any other commonly used unit from
aggregation error; MNI values fluctuate depending on how stratigraphic or cultural units are
grouped, which can have a major effect on how an assemblage is interpreted (Grayson 1984;
Lyman 2008:46). Another problem with MNI is that it tends to overestimate the importance
of rare species in an assemblage. Perhaps the most disconcerting problem with MNI from the
perspective of someone seeking to understand food transport patterns is the assumption that
carcasses, particularly those from large game animals, were brought to a site whole. An
entire body of ethnographic literature (see Chapter 1 for a review) tells us that this is not
necessarily the case. In this study, minimum number of elements (MNE) is preferred for
large mammal biomass calculations, as discussed in Chapter 4. MNI is not used in most of
the analyses below (particularly for large game), though MNI values for each of the species
found at Klissoura Cave 1 are presented in Appendix F.
Minimum number of individuals is used in this chapter for some biomass calculations
(discussed below), because it is a fair assumption that small game animals were brought to
the site in their entirety, provided that the study samples are suitably large. A biomass
comparison allows for a rough estimation of the amount of meat utilized by foragers
occupying Klissoura Cave 1. Often, biomass is calculated based on the mass (weight) of
archaeological bone for each species or group of species in an assemblage (Reitz and Cordier
1983; Reitz, et al. 1987). This method was not employed at Klissoura Cave 1 because the
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lowest Middle Paleolithic layers are heavily mineralized, and bones in certain Upper
Paleolithic layers are covered with thick concretions. As mentioned above, calculations of
small game biomass are based on the assumption that the animals were transported to the site
whole, and are derived by multiplying the average live mass of a given species by the MNI
for that taxon in each stratigraphic layer (Appendix G). Estimated biomass values for large
game species do not rely on the assumption that entire carcasses were brought to the site, and
are calculated by multiplying MNE for each taxon in a stratigraphic layer by a standardized
value of live mass of a given species divided by the number of elements diagnostic to species
level in a given carcass (Appendix G). For example, fallow deer have 55 elements that can be
identified reliably to species (this excludes axial elements and carpal/tarsal complex bones
that may not be recovered archaeologically). The average live weight of an adult fallow deer
is approximately 70 kg, so the standardized value is 70/55 elements, or 1.27 kg per element.
It must be stressed that this biomass calculation is only a rough indicator because it includes
bone and other inedible tissues, ignores variation in individual body size, and assumes that
edible meat is spread evenly over an ungulate body, which it clearly is not. These factors will
be examined in Chapter 6 with a focus on the most commonly occurring species in the
assemblage. For the purposes of this chapter, however, MNE-based biomass estimations can
be used on a broad basis because they standardize NISP for body size and give a more
reliable indication of which species provided the bulk of the meat diet. Species represented
by an NISP<5 for a given stratigraphic layer are excluded from this analysis.
PREY REPRESENTATION IN THE MIDDLE PALEOLITHIC THROUGH
MESOLITHIC LAYERS AT KLISSOURA CAVE 1
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The Klissoura Cave 1 faunal sequence spans much of the end of the Pleistocene, from
about MIS stage 5a until the final deglaciation. This time interval represents variable climatic
conditions that supported a wide range of species near Klissoura Cave 1. Ungulate species
available in Klissoura Gorge, as well as the adjacent upper Berbati Valley and Argive Plain,
included European fallow deer, ibex, European wild ass, aurochs, red deer, and wild pig. Roe
deer and chamois occur in low numbers in Aurignacian layer IV, and roe deer is present
again in the Mesolithic. Small game animals occur throughout the sequence, with tortoises
being common in the Middle Paleolithic and Early Upper Paleolithic. European hare occurs
in low frequencies in the Middle Paleolithic, particularly in layers XI-XIV, but is ubiquitous
in the Upper Paleolithic and later assemblages, when birds such as rock partridge and great
bustard were increasingly important. Large and small carnivore remains are found in low
frequencies throughout the sequence. They include red fox (Vulpes vulpes), wild cat (Felis
silvestris), Eurasian lynx (Lynx lynx), and martens (Martes martes or M. foina). Large
carnivore remains are mainly leopard (Panthera pardus), along with scant remains of wolf
(Canis lupus) in some layers, and a single fragment of brown bear (Ursus arctos) in the
Middle Paleolithic (layer XVIII). Three hyena coprolites were recovered, one in the Upper
Paleolithic, the other two in Middle Paleolithic layers, indicating that hyenas visited the cave,
but their presence was ephemeral. There are absolutely no indications of carnivore dens at
the site. A possible shaft fragment from a hominin (Homo sp.?) ulna was found in the lowest
Middle Paleolithic layer that contained fauna (XXa, XXa1, XXb).
Faunal assemblages from different stratigraphic layers at Klissoura Cave 1 vary in
size. Most notably, the Epigravettian (IIa-d) contains a small sample. The Mesolithic (3-5a),
Early Upper Paleolithic in layer V and Middle Paleolithic layer X are also fairly small. Input
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analysis (Chapter 4) indicates that sample size does not necessarily reflect occupation
intensity, for example the lower Aurignacian (layer IV) and the Middle Paleolithic layers
XVIII-XIX have a similar input rate (measured in NISP count per liter), though the former
has a total sample size of over 4,000 and the latter is around 2,500 (Table 4.1). Tests to
correct for variable sample sizes are undertaken below, and it is still possible to reconstruct
aspects of subsistence from the layers yielding small samples.
Relative species abundance (NISP) through the sequence at Klissoura Cave 1 is
presented in Figure 5.1 (from Appendix F). Several trends are apparent in the data. During
the Middle Paleolithic, fallow deer was the most commonly exploited species at the site.
There is a period of increased prey diversity (evenness) in the Early Upper Paleolithic (V)
and lower Aurignacian (layer IV). Also during the Early Upper Paleolithic (V), small game
becomes a more important part of the diet and increases in frequency until the Epigravettian
(IIa-d) when it actually overtake fallow deer by NISP count. The trends become even clearer
when the Linnaean taxonomic groups are collapsed into the prey categories outlined above.
Figure 5.2 shows the abundance by NISP of the eight prey categories. The medium
ungulate category dominates for the majority of the sequence, though large ungulates attain
greater importance during Aurignacian layer IV. Small, fast-moving game becomes the most
common prey category by count during the Epigravettian (IIa-d) and Mesolithic (3-5a).
Taking this analysis a step further, it is possible to compare changes in the proportion of
large game to small game (combined slow and fast prey categories), as well as changes in the
proportion of small, slow-moving game to small, fast-moving game through the sequence.
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Figure 5.1 NISP counts for the major taxa at Klissoura Cave 1 by layer. Taxa are in descending order
of average mass.
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Figure 5.2 NISP counts for the major taxa at Klissoura Cave 1, grouped by body class.
188
Table 5.2 shows the proportion of small game to ungulate NISP in the Klissoura Cave
1 sequence. The proportion of small game is graphed as a scatter plot against time in Figure
5.3. Warm, mild, and cool phases following the marine oxygen isotope curve (MIS) (see
Chapter 3) are indicated at the top of the graph. As no dates are currently available for the
Middle Paleolithic, points are spaced evenly on the X-axis until MIS 5a, which is thought to
be the stage during which the earliest cultural layers were deposited at the site (Karkanas,
personal communication). Using a Spearman’s rank-order correlation, the relationship
between time period and the proportion of the total assemblage that is composed of small
game is significant and positive (r = 0.754, p = 0.002, n = 14); small game comprise a greater
part of the diet at Klissoura Cave 1 by NISP in the later time periods.
Table 5.2 Proportion of NISP counts per layer for small game: large game (ungulates), and small
slow: small fast-moving prey types. *Small samples excluded from Figure 5.3.
The relative frequency of small game is plotted by period in Figure 5.4, with the two
earliest Middle Paleolithic layers, as well as Middle Paleolithic layer X excluded due to the
small sample sizes for small game (NISP<25). Small, fast-moving animals including the
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Figure 5.3 Proportion of small game to ungulate NISP, plotted against time. The relationship is
significant and positive. Data from Table 5.2.
European hare, rock partridge, and great bustard are depicted using dark shades of grey and
black to indicate their similar handling costs, while tortoises are differentiated by white bars.
Hares were exploited in all periods but their importance increased with time after Early
Upper Paleolithic (V), though they are also common in Middle Paleolithic layer XI-XIV.
Conversely, tortoises were a significant food item only in the Middle Paleolithic and Early
Upper Paleolithic, and are completely absent from the assemblages by the Epigravettian (IIad) (Figure 5.4). Rock partridges and great bustard gained importance in Aurignacian layers
(IIIb-d) and in the period defined by the Mediterranean backed-bladelet industry (III’). The
most productive small prey, hares and partridges, were used heavily by the Upper Paleolithic
and Mesolithic hunters, except during the Early Upper Paleolithic (layer V).
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Figure 5.4 Relative NISP counts for small game animals by layer. White fill indicates small, slowmoving species, grey or black fill indicates small, fast-moving game.
In general, Figure 5.4 indicates an overall shift from small, slow-moving prey species
in the Middle and Early Upper Paleolithic to small, fast-moving animals in the Upper
Paleolithic and later layers. The proportion of small, slow-moving game is graphed as a
scatter plot against time period in Figure 5.5 (data from Table 5.2), again removing the layers
with the small samples mentioned above. A Spearman’s rank-order correlation between time
period and the proportion of the small game fraction that is composed of slow-moving prey is
significant and negative (r = -0.952, p = 0.000, n = 11). Small, slow-moving game animals
were a greater part of the diet at Klissoura Cave 1 by NISP in the earlier time periods, and
were replaced by fast-moving small species in the later part of the sequence. Also, earlier
time periods in general display more variability in the small game animals that were
exploited; by the Aurignacian and later layers tortoises drop out of the assemblages and
small, fast-moving game animals were the exclusive focus.
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Figure 5.5 Proportion of small slow to small fast game NISP, plotted against time. The relationship is
significant and negative. Data from Table 5.2.
Carnivores make up a small proportion of the Klissoura Cave 1 faunal series. Taxa
with an NISP>1 for the entire assemblage are depicted in Figure 5.6. One specimen identified
as brown bear was found in Middle Paleolithic layer XVIII and a single stone or pine marten
specimen was recovered from Aurignacian layer IV (Appendix F). The most common
carnivore species in the series are wild cat, red fox and leopard, though their frequencies are
always low (NISP<25). These taxa may have been utilized for meat and fur, though their
occurrence is low. It is notable that none of the carnivore remains are from young animals, so
it is unlikely that any of the remains are the result of denning behavior (see Chapter 4).
In addition to vertebrate taxa, land snails were exploited during the Upper Paleolithic
at Klissoura Cave 1 (Starkovich and Stiner 2010). No quantitative account of land snails are
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Figure 5.6 NISP counts for carnivore species at Klissoura Cave 1 by cultural layer.
available, but a sample from squares AA4-BB4 was analyzed by M. C. Stiner. The most
common taxon is Helix figulina, a large edible snail found in southeastern Europe. Land snail
shells are not found in the Middle Paleolithic layers, and they are rare in the early Upper
Paleolithic. Shells from the Early Upper Paleolithic and Aurignacian series IV-IIIg show no
evidence of human modification. Shell size in these layers varies, and some shells have small
perforations from predators. The number of land snail shells in the deposits increases with
time. They are moderately abundant in Aurignacian layers IIIb-d and increase greatly in the
Mediterranean backed bladelet industry (III”). Snail shell frequencies are low in the
Epigravettian (IIa-d), like other faunal remains, and peak in the Mesolithic (3-5a). Land snail
shells from the younger layers are biased toward large individuals (2.3-2.4 cm modal
diameter), and display high frequencies of broken aperture rims (75-95%). No snail shells are
burned. It is unclear exactly when land snails start to be commonly exploited at Klissoura
193
Cave 1, though their use is confined to the Upper Paleolithic and later periods. Land snails
are considered a low-return resource because, though they are easy to collect, they are
extremely small and labor intensive to process.
The overall increase in small game exploitation and the specific targeting of small,
fast-moving prey species and small invertebrates such as land snails indicates a greater
reliance on lower-ranked resources in the later layers at Klissoura Cave 1. This may
correspond with human population growth in the region or a more intensive use of the
landscape immediately around Klissoura Cave 1 in the later Upper Paleolithic and Mesolithic
periods. On the other hand, variation in the frequency of great bustards and certain large
ungulates may reflect climate-driven changes in environmental conditions (see below).
PREY DIVERSITY: RICHNESS AND EVENNESS
Taxonomic richness often correlates with sample size (e.g. Grayson 1984; Leonard
1989; Sharp 1990), which simply means that larger faunal assemblages tend to contain more
unique or rare species. In evaluating whether or not this is the case at Klissoura Cave 1,
logNISP is plotted versus logNtaxa (Figure 5.7, values from Table 5.3). Indeed, a Pearson’s
correlation indicates that there is a very significant, positive relationship between the two
variables (r = 0.912, p = 0.000, n = 14). Because sample size explains > 80% of variation,
richness (N-taxa) values at the species level are not useful at Klissoura Cave 1 for
understanding taxonomic diversity in the assemblages because the largest samples contain a
greater number of species.
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Figure 5.7 Ratio of logNISP to logNtaxa in the Klissoura Cave 1 assemblage. The relationship is
significant and positive. Data from Table 5.3.
Taxonomic evenness also sometimes correlates with sample size (Grayson and
Delpech 1998; Grayson, et al. 2001; Nagaoka 2001, 2002b). A Pearson’s correlation between
evenness (1/D of all taxa) and logNISP (all taxa) in Klissoura Cave 1 is not significant (r =
0.165, p = 0.573, n = 14) (data from Table 5.3 and 5.4), which indicates that sample size is
not driving differences in the evenness index. Potential relationships between sample size
and evenness for the ungulate portion of the assemblage as well as the small game
component are also tested; no relationship is found in either of these cases (r = -0.395, p =
0.162, n = 14 for ungulates; r = 0.227, p = 0.503, n = 11 for small game).
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Table 5.3 LogNISP and logN-taxa values for all species, ungulates, and small game species.
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Table 5.4 Inverse Simpson’s index per layer for all taxa, ungulates, and small game species. *Small
sample sizes excluded from Figure 5.10.
In order to evaluate if there are any trends in prey evenness at Klissoura Cave 1, the
reciprocal of Simpson’s Index for all taxa in each layer is plotted against time in Figure 5.8
(Table 5.4). There is a general tendency for an increase in diversity of species exploited from
the Middle to Upper Paleolithic and later periods; this reflects the dominance of fallow deer
in the Middle Paleolithic giving way to greater proportions of small game species at the end
of the Upper Paleolithic. However, the relationship between evenness and time is not
significant (r = 0.477, p = 0.085, n = 14). The outlying point in the Upper Paleolithic
represents Aurignacian layer IV, where there was a more even representation of many largebodied ungulates shown in Figure 5.1. Much of the variability in evenness occurs in MIS 3,
which experienced many climatic fluctuations and shifts in vegetation (see Chapter 3). It is
possible that these rapid shifts are reflected to some degree in the fluctuating diversity of the
Klissoura Cave 1 assemblage.
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Figure 5.8 Inverse Simpson’s index for all taxa plotted against time. No statistically significant
relationship exists, but there is a general increase in variability between assemblages in the Upper
Paleolithic. Data from Table 5.4.
No significant trends in taxonomic evenness are apparent in either the ungulate prey
(r = 0.385, p = 0.175, n = 14) or small game species (r = 0.342, p = 0.304, n = 11) at
Klissoura Cave 1 (Figures 5.9 and 5.10). In general, evenness values are low in the Middle
Paleolithic, while in the Upper Paleolithic and later layers values are variable, which may be
climatically driven, or indicate more flexible foraging strategies. The lower Aurignacian
(IV), Epigravettian (IIa-d) and Mesolithic (3-5a) occupations exhibit greater degrees of
ungulate evenness, which may reflect environmental conditions that will be explored further
below. Small game evenness sorts into two groups, one set with higher diversity, the other
with lower. The three points in Figure 5.10 that are higher diversity likely reflect two
different processes. One of the points represents the Early Upper Paleolithic (Uluzzian layer
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Figure 5.9 Inverse Simpson’s index for all ungulate taxa plotted against time. No statistical
relationship is apparent, though certain Upper Paleolithic and later assemblages are significantly more
even than the other layers. Data from Table 5.4.
V), which has a high proportion of both tortoise and hare. The other points in Figure 5.10
represent Upper Paleolithic layer III” and Aurignacian layer IIIb-d, both of which have
higher frequencies of great bustard and rock partridge (Figure 5.4). The latter set of points is
likely a reflection of drier climates and open vegetation in the vicinity of Klissoura Cave 1.
The declining importance of tortoises, on the other hand, cannot be explained by climatedriven changes in the presence of these species in southern Greece. The decrease in tortoise
numbers in relation to hares corresponds with a shift from high ranked small, slow-moving
species to low ranked small, fast-moving species at the site.
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It is possible that changes in taxonomic evenness are simply a result of changes in site
occupation intensity. For example, more prey species that are rare in the environment may
appear in an assemblage as human hunters spend more time at a site, resulting in a negative
correlation between input and evenness. This is not the case at Klissoura Cave 1, as the
relationship between evenness and input is not significant for all taxa (r = -0.007, p = 0.980,
n = 14). Likewise, there are no relationships between ungulate or small game evenness and
faunal input (r = -0.335, p = 0.241, n = 14 for ungulates, r = 0.184, p = 0.587, n = 11 for
small game). Layers with low evenness values are not necessarily those that have higher
inputs of fauna; rather, evenness may relate to environmental factors or prey choice.
Figure 5.10 Inverse Simpson’s index for small game species plotted against time. No temporal trend
exists, but note the cluster of points that represent higher small game diversity. The earlier point
indicates a more even representation of hares and tortoises, while the later points indicate layers with
an even representation of hares and birds (great bustard and rock partridge). Data from Table 5.4.
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BIOMASS COMPARISON
Analyzing prey biomass provides a method for evaluating the importance of prey
species with different body sizes to forager diets. It goes beyond NISP counts, which are
valuable for understanding changes in prey representation, but does not necessarily account
for difference in the body size of prey being exploited by past hunters, or for differential
fragmentation rates between animals of different sizes (see Chapter 4). For example, the
NISP of hare and fallow deer may each be 50 for a given assemblage. This might result in an
MNI of three hares and one fallow deer (for this example, assume that the animals were
brought to the site whole). Even though the number of hares in the assemblage is greater, the
contribution of fallow deer meat to the diet is greater because the animal is much larger than
even three hares (Table 5.1).
The biomass contribution of each major prey taxon at Klissoura Cave 1 is presented
by layer in Figure 5.11, and is arranged similarly to Figure 5.1. Once again, fallow deer is the
most important ungulate taxon by mass for the majority of the sequence, with the exception
of Aurignacian layer IV, and small game seems to increase in importance by the late Upper
Paleolithic (Figure 5.11). A major difference between the figures, however, is that the
importance of other ungulate species in most of the Middle Paleolithic layers, and some
Upper Paleolithic layers, is much more apparent when biomass is considered (Figure 5.11).
The portion of the diet comprised of small game is much more modest using this method,
even in the Epigravettian (IIa-d) and Mesolithic (3-5a) assemblages that are dominated by
small game NISP counts (Figure 5.11).
Temporal changes in the relative proportion of different groups of species at
Klissoura Cave 1 remain significant when considered in terms of biomass. The proportions of
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Figure 5.11 Estimated biomass for each important taxa at Klissoura Cave 1 by layer. Note that small
game biomass is calculated based on MNI, while large game biomass is calculated using MNE.
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ungulate to small game (all prey types) biomass are listed in Table 5.5. The proportion of
small game biomass is plotted against time in Figure 5.12. A Spearman’s rank-order
correlation indicates that the relationship between small game biomass and time is significant
and positive, excluding the Epigravettian (IIa-d) outlier (r = 0.669, p = 0.012, n = 13).
Changes in the proportion of small game biomass mirror changes in the comparisons based
on NISP, with small game becoming increasingly important through time. Despite the rising
importance of small game in the diet, it must be noted that small game never exceeded so
much as a quarter of the total meat diet by these calculations, with the exception of the
Epigravettian period (IIa-d), where small game comprises more than half of the meat biomass
(Figure 5.12).
Figure 5.12 Proportion of small game to ungulate biomass, plotted against time. The relationship is
significant and positive. Data from Table 5.5.
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Table 5.5 Proportion of biomass values per layer for small game: large game (ungulates), and small
slow: small fast-moving prey types. *Small samples excluded from Figure 5.13.
Considering only the small game component and dividing prey by evasion tactics, it
becomes clear that changes in relative biomass mirror those of NISP counts (Table 5.5,
Figure 5.13). Figure 5.13 again excludes the three Middle Paleolithic layers with small
sample sizes. In general, there was a shift from small, slow-moving species to small, fastmoving species by meat weight through the sequence at Klissoura Cave 1 (Figure 5.13). The
relationship is significant and negative (r = -0.915, p = 0.000, n = 11).
There are subtle differences between the results based on NISP and those based on
biomass, but the larger conclusions are the same. Specifically, changes based on biomass
show the importance of a range of ungulate species in some layers, even if fallow deer are the
most common species as measured by NISP. However, both approaches indicate that across
the Paleolithic sequence at Klissoura Cave 1, small game animals became increasingly
important, and later foragers relied heavily on small, fast-moving game in particular.
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Figure 5.13 Proportion of small slow to small fast game biomass, plotted against time. The
relationship is significant and negative. Data from Table 5.5.
CONCLUSIONS
Shifts in the representation of different prey groups at Klissoura Cave 1 likely signify
changes in species availability as a function of both climate change and hunting pressures
exerted by Pleistocene humans. No trends in species diversity are apparent in the assemblage,
but there is a shift in the proportions of certain types of game through the sequence. One of
the central goals of this presentation is to tease out differences between environmental
impacts on biotic communities and human demographic pressures that may have influenced
the availability of desirable species. It is relevant at this point to return to the predictions
presented in Chapter 3 in an attempt to evaluate whether some shifts in game use at Klissoura
Cave 1 track changing environmental conditions, and if others may be attributed to humans.
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Table 5.6 includes the predictions set forth in Table 3.1, with a simplified summary of
species actually found in different cultural layers at the site. As discussed further below, most
species key to understanding changes in human demography are flexible in terms of their
habitat requirements, and are therefore not useful for environmental reconstructions. This
reduces the risk of tautologies when trying to distinguish between human or environmental
factors as causing changes in the composition of the faunas at Klissoura Cave 1.
The earliest phases of human occupation are thought to have been deposited at
Klissoura Cave 1 as early as MIS 5a, though some layers may also have formed during MIS
4 (XXa-XXb and XVIII-XIX). During milder MIS 5a, wild pigs, fallow deer, tortoises, hares
and partridges should have been available in the vicinity of the site, while during cool, dry
MIS 4, fallow deer, ibex, wild ass, aurochs, tortoise, hare, partridge, and great bustard are
expected. What in fact is found is a dominance of fallow deer, though small amounts of
steppe species including aurochs and European ass, as well as ibex in the oldest Middle
Paleolithic layers, while wild pigs are more abundant in Middle Paleolithic layer XVIII-XIX.
Few small game species were recorded. All available climatic records from this period come
from northern or central Greece, so it is possible that forest expansions recorded elsewhere
during MIS 5a were less widespread in the south, leading to a more mixed environment in
Peloponnese that would have supported the range of ungulate species mentioned above.
Clearly, forests persisted to some degree as evidenced by wild pig, but it seems that these
forests were probably broken up by open areas. Based on the faunal evidence alone, Middle
Paleolithic layers XXa-XXb and XVIII-XIX cannot confidently be assigned specifically to
MIS 5a or 4.
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Table 5.6 Taxa found in Klissoura Cave 1 by cultural layer, with predictions from Table 3.1 in
parentheses. P = present, A = absent, U = uncommon (NISP < 5). Grey shading indicates deviations
from predictions. Roe deer and chamois are excluded because of small samples overall.
*Epigravettian NISP = 67.
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Middle Paleolithic layers XV-XVII, XI-XIV and X either formed toward the end of
MIS 4 or beginning of MIS 3, and layer VIII was deposited during MIS 3. The sequence
lacks dates from the Middle Paleolithic, and it is difficult to distinguish between MIS 4 and
cool periods of MIS 3. Oxygen isotope stage 3 was highly variable and biotic communities
shifted between dry, open periods that would have supported steppe species such as aurochs,
European ass, ibex and great bustard, wet, forested periods in which wild pigs thrived, and
periods of mixed forest-steppe that supported the widest range of prey species: red deer, roe
deer, chamois and ibex. Fallow deer, tortoises, European hare and rock partridges are wellsuited to any of these climatic regimes. Unfortunately much of the climatic variability is
difficult to correlate with the resolution available from the archaeological evidence, and it is
not clear which layers were formed during MIS 4 or 3. In some instances, taxa present at
Klissoura Cave 1 may be indicative of climatic conditions during specific phases of
occupation.
Environmental conditions at Klissoura Cave 1 seem to have been similar for the
duration of the Middle Paleolithic MIS 4/3 (layers XV-XVII, XI-XIV and X) and 3 (VIII).
Fallow deer continue to dominate the assemblages, though steppe species including aurochs
and European ass, and forest-preferring wild pigs are also present in all of the layers. Red
deer and ibex are also found, as well as trace amounts of roe deer. It is unclear if the lack of
great bustard reflects human hunting decisions, or the environment not being sufficiently dry
and open. However, most Middle Paleolithic assemblages in the Mediterranean region lack
human-associated bird remains (but see Blasco and Fernández Peris 2009).
From the composition of species, it seems that most of the Middle Paleolithic layers
were deposited at a time when the Klissoura Gorge and surrounding area was a mosaic of
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forest and steppe communities. The climate may have been on the drier end of the spectrum
to still support mixed plant communities, as it has been established elsewhere that decreased
moisture generally correlates with decreases in primary productivity and a corresponding
decrease in mammalian taxonomic richness (Brown 1973, 1975; Grayson 1998, 2000;
Meserve and Glanz 1978). Richness may not be the most appropriate measure to use at
Klissoura Cave 1 because it correlates so closely with sample size, but evenness values are
quite low during the Middle Paleolithic, which may indicate that diverse ungulate resources
were not usually widely available, especially when conditions were slightly drier.
An increase in moisture is typically followed by an increase in plant productivity,
which is reflected in an increase in the taxonomic richness of animal communities, but only
to a point, since environmental heterogeneity is also important (Abramsky and Rosenzweig
1984; Brown 1973, 1975; Meserve and Glanz 1978). This seems to be the case by the
beginning of the Upper Paleolithic. The Early Upper Paleolithic (V) and lower Aurignacian
(IV) layers display higher levels of species diversity (evenness). Fallow deer and small game
taxa, excluding bustard, are represented in these occupations, alongside species that prefer
steppe communities (aurochs and European ass), forests (wild pigs), and mixed forest-steppe
habitats (red deer, ibex, chamois and roe deer). This range of species, as well as local
climatic reconstructions (Albert 2010; Ntinou 2010), indicates that the plant communities
continued to be mixed forest-steppe, but the taxonomic evenness suggests that conditions
were slightly wetter than they were during the final Middle Paleolithic occupations.
The middle Aurignacian (IIIe-g) and Upper Paleolithic layer III” contain a similar set
of fauna even though they are not consecutive stratigraphically. In many ways, the large
game composition is reminiscent of the MIS 4/3 Middle Paleolithic series, with a dominance
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of fallow deer and low incidence of aurochs, European ass, red deer and ibex. These species,
and the lack of wild pig, indicate that there were probably mixed forest-steppe communities
in the region, and the environment was slightly drier than it was when the first two Upper
Paleolithic layers were deposited. Ungulate species evenness is also low in these periods, like
the later Middle Paleolithic layers. These Upper Paleolithic layers differ from the Middle
Paleolithic, however, because there is a higher proportion of small game.
The upper Aurignacian (IIIb-d) is stratigraphically above the middle Aurignacian
(IIIe-g) and below the non-Aurignacian Upper Paleolithic industry (III”), but its fauna has
more in common with layer III’, the Mediterranean backed-bladelet industry that overlays the
non-Aurignacian Upper Paleolithic (III”). The large game signature is similar to the Upper
Paleolithic layers discussed previously, with an abundance of fallow deer and modest
remains of other ungulate species that prefer steppe or mixed habitats. These two layers are
unique, however, because they contain the highest proportion of great bustard remains. There
is little reason to believe that Pleistocene humans had ignored great bustard earlier in the
Upper Paleolithic; other small, fast-moving species such as hares are found throughout the
sequence. It seems likely that the difference was in prey availability in the gorge area; the
upper Aurignacian (IIIb-d) and the Mediterranean backed-bladelet industry (III’) may have
been deposited during a period of climatic deterioration, when there was a further expansion
of open, steppe communities. These are also the layers in which rock partridge is the most
abundant, though partridges were exploited in other Upper Paleolithic periods. Perhaps
partridges were caught with the same technologies used to hunt bustards and they were
selected as part of an optimal foraging strategy. It is likely that rock partridges flourished
during these periods near the site because, though they are tolerant of a wide range of
210
habitats, they prefer open areas for breeding. A major difference between the two layers is
the increased abundance of hares, a versatile species, in the Mediterranean backed-bladelet
industry (III’) layer, which cannot readily be explained by environmental factors.
Local climatic data that coincide with the Epigravettian (IIa-d) occupation of
Klissoura Cave 1 indicate dry conditions at this time (Albert 2010; Geraga, et al. 2005;
Karkanas 2010; Ntinou 2010). Significantly, there is an absence of great bustard from this
layer. Ungulate evenness is comparatively high, probably because fallow deer frequencies
decline relative to other prey types. Steppe taxa and those that require some forests are
apparent, and their presence indicates that even if the climate was dry, some mixed
communities persisted.
The Mesolithic (3-5a) was deposited during the early Holocene, which was a time of
increased moisture across much of Greece. The species representation in Klissoura Cave 1
does not necessarily reflect this; the same suite of species found in earlier times of mixed
environments are still apparent, and in fact wild pigs are absent from the assemblage.
Conditions may have been similar in MIS 5a, when expanded forests were expected, but are
not actually reflected in the local ecosystems of southern Greece. The Mesolithic (3-5a)
faunal assemblage at Klissoura Cave 1 is similar to that found in the Epigravettian (IIa-d),
with a more even representation of ungulates driven by the decrease in fallow deer, and an
increased importance of European hare.
From an environmental perspective, the major changes that stand out in the Klissoura
Cave 1 faunal series are an increasingly wet environment during the early phases of the
Upper Paleolithic, and a few severely dry periods within the Aurignacian and later industries.
Other trends in game use are more readily explained by human hunting pressures. A major
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shift in the large game component at Klissoura Cave 1 that cannot be readily explained by
climatic change is the relative decrease in the proportion of high-return ungulates to lowerreturn small animal taxa through the sequence, which bottoms-out in the Epigravettian (IIa-d)
and Mesolithic (3-5a). This change is exemplified mainly by a decrease in fallow deer, a
species that dominates the Middle and Upper Paleolithic assemblages by NISP and biomass
until the Epigravettian (IIa-d), when it is surpassed by hares and other ungulate species.
Chapter 6 explores this shift in more detail, considering, for example, whether people were
travelling further for ungulate resources, or if younger animals were exploited more in later
periods. Both of these possibilities may be indications of resource stress.
Another important area of inquiry concerning Paleolithic human diets is changes in
small game use through time (Munro 1999, 2004; Stiner 2001, 2005, 2009; Stiner, et al.
2000). At Klissoura Cave 1, small game use changed in fundamental ways over the course of
the Paleolithic. In addition to a trend towards greater exploitation of small game animals at
Klissoura Cave 1, it has been shown elsewhere that targeting specific kinds of small game
taxa is a sensitive indicator of intensification efforts (Munro 1999, 2004; Stiner 2001; Stiner,
et al. 2000). Specifically, a shift from easy-to-catch small, slow-moving (i.e., higher return)
animals to small fast-moving (i.e., low return) prey is often indicative of increased human
population pressures. During the Middle Paleolithic at Klissoura Cave 1, people relied more
heavily on tortoises to supplement their diet, even though it was always incidental to large
game hunting in terms of absolute meat mass. The proportion of tortoises in the small game
assemblages slowly declines until they disappear from the record after the Mediterranean
backed-bladelet industry (III’). Interestingly, great bustards also disappear from the sequence
at this time, even though the Epigravettian climate was dry and open, conditions preferred by
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this species. A commonality between tortoise and bustards is their low reproductive rates, so
it is possible that human population densities reached a point in these later periods that such
vulnerable taxa were essentially hunted out in the vicinity of Klissoura Gorge. Conversely,
the frequency of hares, which are prolific but low-return, increase through the sequence. It
should be noted that hares are common in one of the Middle Paleolithic layers (XI-XIV) by
NISP count, though they only represent two individuals. It is possible that hares were
occasionally hunted by Middle Paleolithic hominins, or they may be intrusive into these
levels (see Chapter 6 for a discussion of low frequencies of hare burning as compared to
other prey groups in the layer). Along with an increase in small, fast-moving game there is an
increased use of low-ranking land snails in the later Paleolithic and Mesolithic layers. Snail
exploitation began in the later Aurignacian or during one of the Upper Paleolithic
blade/bladelet industries and intensified in later periods.
The Klissoura Cave 1 faunal assemblages provide a long sequence over which
climate change and human hunting decisions can be evaluated. Ideally, many local highresolution environmental studies would be available, in addition to a tight series of dates
spanning the entire archaeological sequence. The available data, particularly from the
Klissoura Cave 1 Middle Paleolithic, currently fall short of this. Despite this shortcoming, the
faunal series at Klissoura Cave 1 can reasonably correlated to larger climatic shifts that are
well-documented across Greece and the Mediterranean Basin, specifically climatic
amelioration during Aurignacian layer IV and a dry, open environment during the
Epigravettian (IIa-d). From the faunal sequence, a convincing case can be made for a more
intensive use of animal resources by human hunters utilizing Klissoura Cave 1 over the
course of the Paleolithic. Multiple lines of evidence, including changes in the frequency of
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ungulate taxa, in particular fallow deer, shifts from high-ranked to low-ranked small game
species, and the disappearance of taxa sensitive to disruptions in their reproduction all point
to the same conclusion: human population densities in southern Greece were on the rise in
the Paleolithic to the extent that new foraging strategies and intensification efforts were
adopted to keep up with this growth. At the same time, biotic communities were forever
altered by human hunters that would turn to agricultural lifeways in a few short millennia.
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CHAPTER 6: PREY SELECTION, TRANSPORT DECISIONS
AND BUTCHERY DAMAGE
INTRODUCTION
Many decisions accompany the subsistence pursuits of hunter-gatherers, from the
location selected for foraging to the animal targeted in a herd. Once a prey animal is
captured, the meat is divided and often transported back to a base camp or habitation site. At
camp, the carcass is processed, which may involve some combination of defleshing, bone
marrow extraction, cooking, and bone grease rendering. All of these behaviors play a part in
determining the composition of faunal assemblages. The series of decisions that goes into
killing and processing an animal may be influenced by many factors, including the size of the
animal, the distance from the base camp where the kill occurs, how many members are in the
group, and how badly food is needed (see Binford 1978; Bunn, et al. 1988; O'Connell, et al.
1988; 1990 for ethnographic examples). I return in this presentation to patch choice and
central place foraging models that were first discussed in Chapter 1; such models are useful
for evaluating transport decisions and butchery patterns across a carcass.
Diachronic change in food collecting behaviors may indicate intensification efforts,
which are often related to resource stress. Intensification can take many forms: the selection
of lower-ranked prey species (Chapter 5), hunting lower-return individuals within a species
(e.g., juveniles) (Broughton 2002; Butler 2001; Davis 1987; Munro 2004; Speth and Clark
2006; Stiner, et al. 2000), hunting forays further from the base camp (Bird and Bliege Bird
1997; Kelly 1995; Metcalfe and Barlow 1992; O'Connell, et al. 1988, 1990), and intensified
carcass processing such as grease rendering (Binford 1978, 1984b; Broughton 1999; Burger,
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et al. 2005; Lupo and Schmitt 1997; Manne and Bicho 2009; Munro and Bar-Oz 2005; Potter
1995; Saint-Germain 1997; Stiner 2003b).
Findings reported in Chapter 4 make it clear that the Paleolithic and Mesolithic
faunas at Klissoura Cave 1 were accumulated and subsequently modified primarily, if not
exclusively, by humans. This chapter explores human activities in greater detail, by
examining the degree of burning on faunal remains, butchery damage, transport decisions
and prey mortality patterns. Large and small game data are evaluated separately because
different considerations must be taken into account in the analyses of transport and butchery
as a function of prey body size. For example, small animals are easily transported whole to a
site by one person (Bartram, et al. 1991; Yellen 1991), whereas larger animals require a
greater effort and are often butchered in the field to maximize the edible meat that can be
carried to the base camp by a small group of hunters (e.g., Binford 1978; Bunn, et al. 1988;
O'Connell, et al. 1988, 1990). This chapter examines two major stages of human decisionmaking in meat procurement and processing: decisions that hunters make in the field in
regards to prey choice and field processing at or near kill sites, and butchery strategies that
are applied once a carcass has been brought to the residential site. Kill and residential sites
may contain a complementary set of activities, and only the residential site is represented at
Klissoura Cave 1. A single large skeleton may be divided between a kill site and home base,
with bulky or low-utility elements abandoned at the kill, and meaty or easily carried elements
transported to a residential site. Evidence of defleshing or disarticulation may begin at the
kill site but continue at a base camp. Though some snacking and cooking can occur at kill
sites, burning, intensive marrow processing, and grease rendering are typically activities that
are confined mainly to residential sites. Based on the large number of hearths, the
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Aurignacian layer IV structure, the location of the site in a rock shelter, and many other
indications, it is reasonable to assume that Klissoura Cave 1 was used primarily or
exclusively as a base camp or residential site. Therefore, it is expected that the
zooarchaeological assemblage should reveal processing activities that occurred at the
residential site, and the suite of skeletal elements that were selected for transport back to the
shelter. Both lines of evidence are useful indicators for understanding whether hominins at
Klissoura Cave 1 were making decisions based on optimizing their returns or intensively
harvesting food resources.
Many observations associated with human activities were recorded during the
analysis of the Klissoura Cave 1 faunas, including burning damage, tool marks, impact
fractures, and fresh bone breaks. Damage on faunal specimens of both large and small prey
species confirms that the faunas were collected and modified by humans (Chapter 4), a point
that is expanded upon below. In the analyses presented in this chapter, NISP counts often
vary between tables and with values presented elsewhere in this dissertation, following points
made in Chapter 4; shattered dental elements or certain bones that are highly recognizable
because of their texture (e.g. tortoise shell fragments) are used in taphonomic analyses, such
as burning frequency, but not in considerations of species representation. Some of the tests
presented below, such as those that evaluate butchery damage, exclude dental specimens
because their fragmentation often is not directly related to food processing. Other tests target
specific anatomical elements, as noted in the sections that follow.
BURNING DAMAGE
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Burning stages on the faunal remains were recorded following Stiner and colleagues
(2005:45; 1995:226) (Table 6.1). Seven stages were recorded, including unburned, three
different degrees of carbonization (semi- to fully blackened), and three different degrees of
calcination (semi- to fully whitened). Browning of bones was not recorded as burning, since
many other factors, such as staining, can produce coloration in archaeological bone.
Published bone burning experiments indicate that fires using hardwoods common in the
Mediterranean easily reach a temperature of 900 °C (Stiner, et al. 1995). Bone becomes
calcined only when it is in direct contact with fires at these temperatures (Stiner, et al.
2005:50; Stiner, et al. 1995:234). Burning damage does not necessarily indicate that meatcovered bones were cooked to the point of charring, or even that bones were deliberately
thrown into a fire after a meal. Experimental studies indicate that hearths built on sediments
containing bone fragments burn bones up to 5 cm below the surface, depending on the
sedimentary matrix and duration of burning, though these specimens rarely reach the point of
calcination (Bennet 1999; Stiner, et al. 2005:50; Stiner, et al. 1995:230). This means that
archaeological faunas can be carbonized long after they are deposited, though the highest
levels of burning must come from direct contact with fires and may concentrate in the
vicinity of hearth areas in sites that are visited repeatedly. There is certainly the potential for
post-depositional burning at Klissoura Cave 1, with the abundance of hearth structures
through the layers (Karkanas, et al. 2004; Koumouzelis, Ginter, et al. 2001; Pawlikowski, et
al. 2000).
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Table 6.1 Burn frequency and degree for different classes of animals at Klissoura Cave 1 by layer.
219
Table 6.1, cont. Burn frequency and degree for different classes of animals at Klissoura Cave 1 by
layer.
220
Table 6.1, cont. Burn frequency and degree for different classes of animals at Klissoura Cave 1 by
layer.
221
Bone is also useful as a fuel source in some circumstances; experimental studies
indicate that it extends the life of a fire depending on how much bone is added (Théry-Parisot
2002), though the addition of small fragments tends to smother a fire (Mentzer 2009).
Experimental studies using bone as fuel yields assemblages with high frequencies of calcined
bone (76.1%) and high rates of fragmentation (56.9% smaller than 2 cm), particularly of
cancellous bone (Cain 2005; Costamagno, et al. 1998; Costamagno, et al. 2005; ThéryParisot 2002). Establishing a connection between experimental bone burning studies and
archaeological evidence of bone as a fuel source is difficult, however, because calcined bone
tends to preserve poorly in archaeological contexts (Costamagno, et al. 2005; Stiner, et al.
1995), and cancellous bone is more susceptible to density-mediated processes (see Chapter
4). Mineralogical studies of archaeological sediments may help bridge the gap between
experimental work and the actual use of bone as fuel in past with the identification of
microscopic ash or calcined bone (Mentzer 2009).
Burning frequencies are presented for different categories of species (i.e., tortoise,
medium ungulate, etc.), as well as for each layer as a whole to determine if certain species
categories are burned more often than others. This also helps to identify potentially intrusive
animals, such as carnivores. Tooth and bone elements are included in this comparison. An
association between degree of fragmentation and burning is well-established at many
archaeological sites (Cain 2005; Costamagno, et al. 1998; Costamagno, et al. 2005; Stiner
2005:96-97; Stiner, et al. 1995:224; Théry-Parisot 2002). Mean fragment length for burned
and unburned specimens at Klissoura Cave 1 is presented to determine if there is a similar
difference at the site, particularly in relation to hearth areas (see also Chapter 7), and to
evaluate the possibility of burning bone for fuel. Burn damage within species was also
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examined to see if certain elements were treated differently. Tortoise remains in the Hayonim
Cave and Meged Rockshelter assemblages of Israel, for example, show differences in
burning frequencies for plastrons (flat, belly plate) versus carapaces (dorsal shell) (Stiner
2005:99), and significant differences in burning rates are found from marrow rich bones at
Kebara Cave (Speth and Clark 2006) and at Qesem Cave (Stiner, et al. 2009). Differences
between the degree of burning for medium ungulate crania and postcrania is also examined.
Teeth are excluded from this particular burning analysis because they tend to shatter when
burned, which would bias the results.
Burning Damage on Small Game
Small game burning damage is presented in Table 6.1. Trends in burning are difficult
to discern for some species because the sample sizes are small. This is the case with tortoise
remains for most of the Upper Paleolithic and Mesolithic layers. In the early Upper
Paleolithic and Middle Paleolithic layers, tortoise remains are heavily burned, often at nearequal or higher rates than the assemblage as a whole (with the exception of Middle
Paleolithic layers VIII, XI-XIV and XXa-XXb, the latter of which is a fairly small sample)
(Table 6.1). The extent of burning of tortoise carapace and plastron fragments is presented in
Table 6.2. In slightly more than half of the layers with samples of shell fragments larger than
NISP = 20, plastron fragments exhibit equal or higher rates of burning than carapace
specimens. Differences in the frequency of carapace and plastron burning is not as striking as
that observed at some sites in Israel (Stiner 2005), though smaller tortoise shell samples in
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Table 6.2 Burn frequencies for tortoise shells in layers with larger sample sizes.
most layers may explain the discrepancy. High rates of burning on tortoise specimens is
consistent with results obtained from Kebara and Hayonim Caves in Israel (Speth and
Tchernov 2002; Stiner 2005).
Upper Paleolithic and Mesolithic hare specimens exhibit about the same frequency of
burning as the rest of the animal remains in each layer, while Middle Paleolithic hare remains
are burned much less often than their respective whole assemblages (Table 6.1). Hare
remains are fairly uncommon in the Middle Paleolithic layers at Klissoura Cave 1, with the
exception of layer XI-XIV. The fact that hare specimens in all Middle Paleolithic layers are
less burned than the bones of other animals suggests that they may have been intrusive into
the layer, though a lack of avian or mammalian carnivore damage on hare remains (Chapter
4) indicates that they were not introduced to the site by predators.
In layers that contain the largest bird assemblages (Mediterranean backed bladelet industry
III’ and Aurignacian layers IIIb-d and IV), birds have considerably lower rates of burning
than the average rate for these layers (Table 6.1). Carnivores also have extremely low
incidences of burning, with the exception of Aurignacian layer IV, which may be explained
224
by sample sizes (Table 6.1). This may indicate that at least some of the remains were
intrusive, or that they were cooked or disposed of differently than ungulates.
Burning Damage on Ungulates
For the Klissoura I faunal assemblage as a whole there is a statistically significant
trend toward a decrease in burned bone over time (Table 6.1; n = 14, r s = 0.556, p = 0.039).
This is reflected in the degree of burning for ungulates, which comprise the bulk of the
assemblage through most of the sequence. Burning rates are similar for medium and large
ungulates, except when samples for large ungulates are small (Table 6.1). Calcined bones
were recorded in nearly all layers, indicating that some specimens were in direct contact with
fires. The average lengths of burned and unburned specimens are presented in Table 6.3;
burned specimens are consistently smaller than their unburned counterparts in every layer
except the Epigravettian (IIa-d), which has an extremely small sample.
Table 6.3 Mean length for burned and unburned identifiable specimens by layer.
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Variation exists in the frequency of burning damage through the stratigraphic
sequence, with the Middle Paleolithic and Aurignacian layers containing more burned bone
than the later layers. This variation probably relates to occupation intensity as opposed to
specific cooking behaviors; high frequencies of burning are common in layers that contain
large numbers of hearth features (see Chapter 7), though burn frequency does not correlate
with faunal input rates (n = 14, rp = 0.047, p = 0.874).
Table 6.4 indicates the frequency of burning for medium ungulate crania versus postcranial
specimens by layer; the differences are plotted in Figure 6.1 in layers with large enough
sample sizes. Antlers are excluded from this analysis in case they were collected after they
were shed, a strong possibility given the importance of antler for osseous tools. In most of the
Upper Paleolithic layers, cranial elements are burned more often than postcranial elements
(with the exception of Aurignacian layer IIIe-g). No such difference is apparent in the Middle
Paleolithic; in general postcranial specimens have higher rates of burning, though burn
frequencies are fairly close between the two sets of body parts. Differences in burning in
Upper Paleolithic layers may indicate the systematic roasting of medium ungulate crania.
There is little evidence to indicate that bone was used as a fuel source at Klissoura
Cave 1. The average fragment size of burned bone is smaller than unburned bone, which is
the case at other Paleolithic sites (e.g., Stiner 2005:92). The mean fragment size at Klissoura
Cave 1 is larger than 2 cm in all layers except for one, however, and generally greater than
expected if bone was burned for fuel (Cain 2005; Costamagno, et al. 1998; Costamagno, et
al. 2005; Théry-Parisot 2002). There is no overabundance of burned cancellous bone in
Klissoura Cave 1, especially in the Middle Paleolithic, though this may be the result of
226
Table 6.4 Burn frequency and degree for cranial and postcranial elements by layer for medium
ungulates. *Small samples not included in Figure 6.1.
227
Table 6.4, cont. Burn frequency and degree for cranial and postcranial elements by layer for medium
ungulates. *Small samples not included in Figure 6.1.
228
Figure 6.1 Frequencies of burned cranial and postcranial specimens at Klissoura Cave 1. In the Upper
Paleolithic layers crania are more commonly burned.
density-mediated attrition. Calcined bone is expected if bone is used as a fuel source
(Costamagno, et al. 1998), but it is present in low frequencies in all layers. Costamagno et al.
(2005) stress that the unidentifiable component of an assemblage is as important for
evaluating bone burning as the identifiable portion, but calcined bone is also highly
susceptible to degradation (Stiner, et al. 1995). More significant in the case of Klissoura
Cave 1 is that fact that burned bones do not correlate spatially with the presence of hearth
features (see Chapter 7), or in discrete areas that may indicate fireplace cleaning events. At
this stage, there is no evidence that bone was burned as fuel at Klissoura Cave 1.
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BUTCHERY DAMAGE AND OSSEOUS TOOLS: CUTS, CONES, IMPACTS,
WORKING, AND OTHER FRACTURES
Evidence of butchery is perhaps a more direct indicator of human modification of a
faunal assemblage than burning, since it is not always clear if burning damage is associated
with cooking activities, site cleaning, disposal of bone, or if burning occurred postdepositionally. Several kinds of butchery damage were recorded in the Klissoura Cave 1
faunas. Cone, or hertzian fractures and impacts are typically associated with the extraction of
marrow from a mammalian long bone cavity (Binford 1978; Blumenschine and Selvaggio
1991:153-156; Brain 1981:141-142; Stiner 1994) or breaking open a tortoise shell for meat
(e.g. Stiner 1994). Such fractures are occasionally caused by carnivore chewing or crushing,
but are more often associated with human butchery activities, specifically the striking of
cortical bone shafts with a hammerstone. Large carnivores occasionally create cone fractures,
but Paleolithic humans produce them at much higher frequencies (Stiner 2005:93). Tool
marks were recorded in the assemblage, though observed frequencies in the Upper
Paleolithic and later assemblages likely underestimate the true number of cuts because many
specimens were partly or wholly covered with concretions of ash (see Chapter 4). In this
case, the presence of cone fractures or impacts and cut marks are more instructive than raw
frequencies of such damage in establishing that humans modified faunal remains.
Green bone fractures or "fresh breaks" occur postmortem, but while bone still
maintains its natural fibrous composite structure (Currey 1984; Johnson 1985:160). Such
breaks are indicative of damage shortly after the death of an animal, before the bone dries
(Davis 1985:66-67). Many different fracture types are observed on archaeological
assemblages (see Davis 1985; Gifford-Gonzalez 1989:188; Lyman 1994:318-324; Marshall
230
1989; Shipman, et al. 1981). Here I focus on fresh spiral fractures on tortoise bones and
transverse fractures on mammalian and bird bones. In this case, spiral fractures are classic
green bone breaks. Transverse green fractures are similar, but occur when a bone is broken
transversely to its main axis using a shearing action (Gifford-Gonzalez 1989:188; Stiner
2005:93). This type of fracturing may relate to the partitioning of carcasses by hominins
during butchery for transport or sharing (Stiner 1994). Carnivores produce green bone
fractures in much the same way that they cause cone fractures. Some investigators have also
linked transverse fractures on small mammals to human processes, such as on the long bones
of hares and rabbits during marrow removal (Hockett 1994); transverse fractures on small
animal bones may also relate to dismemberment (Stiner 2005:95). In the results presented
below, butchery activities on ungulate remains are from all boney skeletal elements.
Frequencies of cut marks on small animals are likewise counted on all bone elements, while
transverse fractures on small game are only evaluated on selected long bones, including
femora, humeri, tibiae (or bird tibiotarsi) and radii. Spiral fractures are presented for all
tortoise shell fragments in the assemblages.
Butchery Damage on Small Mammals and Carnivores
Frequencies of cut marks on small animal and carnivore remains are presented in
Table 6.5. Cut marks are present but rare on all species throughout the assemblages. The low
frequencies of tool marks may be a combination of small sample sizes for small animals and
carnivores in most layers combined with surface encrustations on specimens, especially in
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Table 6.5 Butchery damage on small game at Klissoura Cave 1 by layer.
the Upper Paleolithic layers (see Chapter 4). For example, the largest samples of hare and
bird bones are from the Upper Paleolithic and later layers, where ashy concretions were
common on the remains and frequencies of cut marks are extremely low (Table 6.5). One cut
marked specimen was recorded for hares in the Middle Paleolithic, in layer XXa-XXb. A few
tortoise elements exhibit cut or impact marks (Figure 6.2), but again frequencies are quite
low.
Evidence of human modifications on small animal remains comes mostly from high
frequencies of transverse fractures on mammal and bird long bones and spiral fractures on
tortoise shells (Table 6.6). Transverse fractures occur on well over 50% of hare long bones
(with the exception of Middle Paleolithic layer XI-XIV) and 60% of bird long bones in
assemblages with sample sizes greater than ten (Table 6.6; see also Bocheński and Tomek
2010 for a further discussion of damage on bird remains). Transverse fractures are also
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observed on carnivore remains, though samples are small for any given layer (NISP < 5).
Humans traces are the only significant modifications of the small animal remains, indicating
that humans were the primary collectors of small game remains at Klissoura Cave 1. Low
frequencies of fractures on Middle Paleolithic hare remains supports the hypothesis that hare
remains were intrusive during much of this time period.
Figure 6.2 Tortoise carapace fragment with crushing and an impact fracture. From Middle Paleolithic
layer XVI.
Butchery damage is found on all groups of small animals at Klissoura Cave 1,
including the small mammals, reptiles, birds and carnivores in the assemblage. Cut marks are
rare on small game remains, which is partially explained by poor bone surface visibility on
Upper Paleolithic and later layers (Chapter 4). Another possibility for hares in particular is
that elements expected to have cut marks associated with skinning (e.g., feet) are generally
lacking in the assemblage (see below).
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Table 6.6 Transverse fractures on small animal limb bones (humerus, radius, femur and
tibia/tibiotarsus) by layer.
Butchery Damage on Ungulates
Butchery damage summaries for medium and large ungulates are presented in Table
6.7. Conservative criteria are used to evaluate damage, so cone and impact fractures are less
common in the Klissoura Cave 1 assemblages than in other Mediterranean Paleolithic sites
(see Stiner 2005:107), occurring on 1-3% of medium ungulate remains in most layers (Table
6.7). Impact damage is observed on a slightly higher proportion of large ungulate specimens
in layers with sample sizes larger than n = 20, at rates of 1-4.5% (Table 6.7). There is no
trend in the frequency of impact damage on either the medium or large ungulate remains (n =
14, rs = -0.525, p = 0.054 and n = 11, rs = -0.301, p = 0.368, respectively), though the p-value
for medium ungulates is nearly significant, with a decrease in impact marks in later periods.
This correlation, to the extent that it indicates a significant relationship between medium
ungulate remains and time period, has the potential to explain 28% of the variation in the
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Table 6.7 Butchery damage on medium and large ungulate remains. *Note small sample sizes.
assemblages, which may reflect small samples in the youngest layers, or be related to
decreased visibility of bone surfaces in Upper Paleolithic layers. In any case, butchery
damage does not increase through time, which indicates that bone marrow was not
necessarily more intensively accessed in later time periods.
Frequencies of tool marks on the Upper Paleolithic assemblages are difficult to assess
because of ashy concretions on the surfaces of bone in these layers. Cut marks are found on
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0.3-1.1% of Upper Paleolithic medium ungulate remains and are almost entirely absent on
large ungulate specimens (Table 6.7). Because tool mark frequencies for the Upper
Paleolithic layers are unreliable due to low visibility of bone surfaces, no further analyses
were attempted on them. Tool marks are more common in the Middle Paleolithic layers on
both medium and large ungulate remains; cut marks are present on 0.8-3.4% of medium
ungulate specimens and 1.1-6.4% of large ungulate remains in samples larger than NISP = 30
(Table 6.7) (see Figure 6.3 for an example of cuts on a medium ungulate specimen). Tool
mark counts for medium ungulates are listed by element in Table 6.8 by layer. Most cut
marks were recorded on unidentified long bone fragments. Aggregated cut frequencies for
the entire Middle Paleolithic are illustrated in Figure 6.4 for elements with a combined NISP
> 10 cuts. Most of the tool marks occur on the hind limbs, mandible, humerus or ribs.
Transverse fractures are common in the ungulate assemblages at Klissoura Cave 1,
with 12-46% of medium and large ungulate remains exhibiting such damage (Table 6.7).
There is a statistically significant trend in the frequency of transverse fractures on ungulate
specimens, with fewer fractures in the older layers (n = 14, rs = -0.895, p = 0.000 for medium
ungulates and n = 11, rs = -0.664, p = 0.026 for large ungulates). This may indicate the
development of finer carcass partitioning for transport, sharing, cooking, or marrow
processing strategies in the later periods.
Worked Bone and Antler
The use of bone and antler as raw materials for tool manufacture is rare in Europe and
western Asia during the Middle Paleolithic (Villa and d'Errico 2001), though these
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Figure 6.3 Example of cut marks on medium ungulate remains. Specimen from Middle Paleolithic
layer XII.
Table 6.8 Cut marks on medium ungulate elements in Middle Paleolithic layers.
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Figure 6.4 Cut frequencies on fallow deer elements in the Middle Paleolithic layers at Klissoura Cave
1. Includes elements with n > 10 cuts.
technologies appear by 80,000 to 100,000 years ago in Africa (McBrearty and Brooks 2000).
The use of osseous technologies in Europe is associated first with basal Upper Paleolithic and
transitional technologies, such as the Châtelperronian, Uluzzian and Streletskaian, among
others. Osseous technology becomes ubiquitous during the Aurignacian (Bar-Yosef, et al.
2006; Conard and Bolus 2003; d'Errico, et al. 2003; d'Errico and Laroulandie 2000; d'Errico,
et al. 1998). Worked bone is present in the Klissoura Cave 1 assemblages, but only in the
Upper Paleolithic and later deposits (Koumouzelis, Ginter, et al. 2001). Nearly all of the
osseous tools and manufacturing debris were separated from the faunal remains as the time of
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this study. Simple counts of worked bone and antler by layer are included in this chapter,
though a more in-depth study of osseous tools from a technological standpoint is ongoing by
Rozalia Christidou.
Overall, worked bone or antler is rare in the assemblages (Table 6.7). One possible
worked bone was recorded in Middle Paleolithic (XI-XIV) (Figure 6.5); the rest of the
worked bone come from Upper Paleolithic and later layers. Most of the osseous technology
at Klissoura Cave 1 is made from antler, so proportions of antler relative to total ungulate
NISP counts are compared in Table 6.9 along with percentages of worked antler by layer.
Antler is present in appreciable frequencies, given that only adult male cervids develop
antlers, and they are present on the animal only on a seasonal basis. Tools made of antler
were found primarily in the middle and lower Aurignacian layers (IIIe-g and IV), though a
few isolated tools were also found in later layers (Table 6.9). Aurignacian layers IIIe-g and
IV also contain high frequencies of antler fragments, suggesting some on-site production of
osseous tools. The non-Aurignacian Upper Paleolithic layer (III”) has a high proportion of
Figure 6.5 Worked bone from Middle Paleolithic layer XIV.
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Table 6.9 Counts of all antler and worked antler by layer, and corresponding ungulate NISP counts
for comparison.
antler specimens, though none of it is worked (Table 6.9). In general, antler is rare in the
Middle Paleolithic layers, indicating that male fallow deer were infrequently targeted for
hunting or that male deer were hunted in seasons when antler was unformed or in the early
stages of growth.
BONE PROCESSING
One measure of intensification of meat resources is the extent to which a carcass is
processed for its edible tissues (e.g., Binford 1978, 1984b; Broughton 1999; Burger, et al.
2005; Lupo and Schmitt 1997; Manne and Bicho 2009; Munro and Bar-Oz 2005; Potter
1995; Saint-Germain 1997; Stiner 2003b). As discussed in Chapter 1, the patch choice model
predicts that the highest-return tissues are utilized first. Lower-utility animal products are
typically only exploited if their processing costs are lower than travel and handling costs for
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additional prey items. Heavy exploitation of low-return tissues may reflect resource stress or
intensification. Organs and muscle are the easiest tissues to extract and require a fairly
minimal degree of butchery. Cold marrow processing is slightly more labor intensive but still
quite simple; using a hammerstone, marrow-rich long bones are crushed, granting access to
nutrient-filled bone marrow cavities, and creating cone and impact fractures discussed above
(e.g., Binford 1978; Blumenschine and Selvaggio 1991; Brain 1981:141-142; Stiner 1994,
2005). This level of processing began in the Lower Paleolithic, possibly as early hominins
scavenged ungulate carcasses. Cold marrow processing from medullary cavities is nearly
ubiquitous in Paleolithic sites from all subsequent periods and regions (e.g., Bar-Oz and
Munro 2007; Binford 1984b; Blumenschine and Magrigal 1993; Boyle 2000; Brain 1981;
Costamagno, et al. 2006; Egeland and Byerly 2005; Gaudzinski-Windheuser and Niven
2009, and discussion therein; Hockett and Bicho 2000; Lupo 1998; Miracle 2005; Munro and
Bar-Oz 2005; Niven 2007; Stiner 1994, 2005; 2009, among many others). In order to assess
the degree of cold marrow processing at Klissoura Cave 1, ungulate long bones are ranked by
the volume of their marrow cavities. Large ungulates are excluded from this analysis because
of small sample sizes. MNE values are listed for each element, and the percent of unopened
elements are indicated. It is expected that elements with the greatest amount of bone marrow
tend to be opened more frequently. It is also predicted that when resources are intensified,
perhaps during periods of stress, elements containing smaller amounts of marrow will be
broken at high frequencies.
An even more intensive use of large game carcasses is heat-in-liquid bone grease
rendering, a process that seeks to unlock small pockets of animal fat in the pores of spongy
bone elements. During grease rendering, cancellous parts of elements are pulverized and
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boiled, typically by placing heated stones into water and liberating the oil, which can be
skimmed from the top of the mixture, purified and stored (e.g., Binford 1978; Brink 1997;
Lupo and Schmitt 1997; Saint-Germain 1997; Vehik 1977). Bone grease rendering is
extremely labor intensive (Church and Lyman 2003), and is typically associated with
increased seasonality (Enloe 2003a; West 1996, 1997) or with resource intensification (BarOz and Munro 2005; Manne and Bicho 2009; Munro 2004; Nakazawa, et al. 2009; Stiner
2003b). Notably, bone grease rendering has not been found in the Mediterranean region
earlier than the Gravettian (Manne and Bicho 2009; Manne, et al. 2005). At Klissoura Cave
1, the most likely layers to contain bone grease rendering (e.g., the Epigravettian and
Mesolithic) have such small samples that it is difficult to test for grease rendering. Further,
supporting artifacts typically found associated with grease rendering such as fire cracked
rock or hammerstones and anvils (e.g., Manne and Bicho 2009; Manne, et al. 2005) are not
common at Klissoura Cave 1 (Kaczanowska, et al. 2010).
Ungulate Marrow Processing
Evidence of bone marrow processing is based in part on the percentages of opened
medium ungulate long bone medullary cavities (Table 6.10). No major long bone elements
are complete in any of the layers, indicating that medium ungulates were processed
consistently for medullary marrow in all periods. Some phalanges were recovered intact,
however. These elements contain the least amount of marrow (see marrow utility indices in
Binford 1978). Table 6.11 presents the percentage of unopened phalanges by layer provided
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Table 6.10 Percent of medium ungulate limb bones not opened prior to discard, by layer.
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Table 6.11 Percent of unopened medium ungulate phalanges for layers with MNE values larger than
10.
that the assemblage was relatively large. In general, first phalanges are processed more
heavily than second phalanges, which were in turn opened more frequently than terminal
phalanges (Table 6.11, Figure 6.6). This reflects the decrease in nutritional return that comes
from opening third phalanges, as they contain less marrow than the proximal phalanges. No
temporal trend exists for processing any of the phalanges, though there is an increase in
opened third phalanges during Middle Paleolithic layer XI-XIV and especially during
Aurignacian layer IV. This may indicate a more intensive use of animal carcasses,
particularly during Aurignacian layer IV, which was a period in which occupation intensity
or site use changed .
A few differences in marrow processing strategies were observed in at Klissoura
Cave 1, though marrow was intensively utilized throughout the sequence. There is no
evidence for bone grease rendering, though by the time this strategy is expected to appear in
Eurasia (during the Gravettian or later), there is an overall shift to lower-ranked small game
resources at Klissoura Cave 1 and much less frequent use of the shelter (see Chapter 5). One
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Figure 6.6 Percent of unopened first, second and third phalanges through the layers at Klissoura Cave
1. First phalanges, which contain the highest marrow content, are consistently processed through the
layers. Second phalanges were opened more often in the late Middle Paleolithic and Aurignacian
layers. Terminal phalanges were only intensively processed during the late Middle Paleolithic and
Aurignacian layer IV (data from Table 6.11).
notable trend in butchery strategies is a statistically significant increase in transverse
fractures in later time periods. This could relate to more standardized carcass processing
strategies in the Upper Paleolithic, or at least more efficient segmenting for transport,
cooking, or while extracting bone marrow. Another difference is the increase in processing of
terminal phalanges, an element with an extremely low marrow yield, during Aurignacian
layer IV.
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PREY TRANSPORT
Evaluating the representation of prey body parts at an archaeological site is
instructive in understanding the decisions surrounding the transport of prey to a base camp or
habitation site, assuming post-depositional taphonomic factors did not produce the damage
(see Chapter 4). Central place foraging models are often used to understand transport
decisions involving the movement of specific portions of large-bodied prey (Broughton
1999; Cannon 2003; Nagaoka 2005; O'Connell, et al. 1988, 1990; O'Connell and Marshall
1989; Speth 1991; Speth and Scott 1989). As local resources are depleted, hunters travel
farther from residential sites to procure large game. The further they travel, the more
selective foragers tend to be about which portions of a carcass are carried to the habitation
site. If the meat must be carried over long distances, foragers typically focus on transporting
fewer high-utility skeletal elements, or elements that provide the greatest nutritional return
(Binford 1978; Broughton 1999; Cannon 2003; Nagaoka 2005; Speth 1991; Speth and Scott
1989; Stiner 1991, 1994, 2005). A bias toward high-utility skeletal portions at Klissoura
Cave 1 may indicate increased travel times between the site and hunting location.
Small-bodied prey can easily be carried to a site in whole form (Bartram, et al.
1991:102; Yellen 1991:5). Examining body part profiles for small animals, such as hares or
birds, is valuable because it contributes to our understanding of taphonomic processes at a
site and the possible presence of discrete activities areas within a site. Of course, the
hypothesis that small game animals were transported to the site whole must also be
evaluated.
Utility Indices
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There are many methods for examining prey transport decisions. One is based around
utility indices introduced by Binford (1978). Based on his work with the Nunamiut in Alaska,
Binford (1978) derived a series of indices for caribou (Rangifer tarandus) and dall sheep
(Ovis dalli) in order to rank the utility of different classes of edible tissue (meat, marrow, and
bone grease) by body part for these two species (see Appendix H for values, from Binford
1978). Two combined indices were derived from the three tissue-specific indices mentioned
previously: the general utility index (GUI) considers the combination of meat, marrow and
bone grease associated with different elements or portions of elements, and the modified
general utility index (MGUI) takes into account food-poor articulated elements that tend to
be transported as riders with larger, food-rich elements. GUI and MGUI values both correlate
significantly with meat index values; MGUI also correlates with bone grease index values
(see Table 6.12). Raw index values are standardized to a scale between one and a hundred,
and the normed MGUI values become %MGUI. In order to compare utility indices with
archaeological data, %MAU is constructed by calculating MAU values for each element, and
dividing this by the maximum MAU value in the assemblage: (observed MNE/expected
MNE)/(maximum MAU). Binford (1978:81) then constructed a series of models of transport
strategies based on scatter plots of %MGUI versus %MAU. This analytical model was
quickly applied to archaeological assemblages by other zooarchaeologists (Landals 1990;
Speth 1983; Thomas and Mayer 1983), though many were critical of aspects of the
technique, particularly its application to taxa for which no index values had been derived
(Chase 1985; Jones and Metcalfe 1988; Metcalfe and Jones 1988). It was later realized that
certain relationships between %MGUI and %MAU (specifically the “reverse utility curve”)
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Table 6.12 Pearson’s correlation values between Binford’s (1978) utility indices. * Indicates
significant relationships.
could be caused by post-deposition attritional processes in archaeological assemblages rather
than human transport decisions (Grayson 1988, 1989; Lyman 1985). Additionally, some
utility curves (specifically the gourmet strategy and reverse gourmet strategy) particularly as
conceptualized by Lyman (1994:228) could never occur because so few elements in an
ungulate assemblage have %MGUI values above 80, which is required for these two models
to be realized.
Despite these potential pitfalls, it is useful to examine relationships between utility
indices and %MAU in zooarchaeological assemblages, if density-mediated effects have
already been largely excluded. Density-mediated attrition was not a significant problem for
the ungulate remains in the Upper Paleolithic and later layers at Klissoura Cave 1. On the
other hand, possible density-mediated biases in Middle Paleolithic layers must be kept in
mind during this analysis. Correlations between %MAU and high utility values, particularly
meat, GUI or MGUI may be indicative of transport strategies that focus on high utility
elements. Although these indices include bone grease, this does not seem to have been a
motivating factor for the transport of certain ungulate elements to the site, as there is no
evidence of bone grease rendering. Thus, the main goals in the transport of ungulates would
include meat and medullary marrow.
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Percent MAU values for fallow deer element portions are listed in Appendix I by
layer; corresponding meat, marrow, grease, GUI and MGUI values for reindeer are in
Appendix H (from Binford 1978). Spearman’s rank-order correlation values for %MAU and
the respective utility indices listed above are presented in Table 6.13. Few correlations are
significant, with the exception of marrow index values in Aurignacian layer IV and Middle
Paleolithic layer XXa-XXb (Table 6.13). It is possible that marrow-rich elements were
preferentially transported to the site during these periods for the added nutritional value of
bone marrow, particularly during Aurignacian layer IV when there are other markers of
intensification (e.g., marrow processing ungulate terminal phalanges). On the other hand,
there is clearly no evidence for preferential transport of elements with overall high utility.
This is especially the case in Middle Paleolithic layer VIII, where there is a significant,
negative relationship between meat and GUI index values and %MAU, indicating a lack of
high-utility elements at the site during this period. It must be noted that several lines of
evidence indicate that density-mediated attrition was a factor in layer VIII (Chapter 4).
Scatter plots of %MAU and %MGUI are presented by layer in Figure 6.7. For the
most part, the plots do not resemble any of Binford’s (1978) models of transport strategies.
However, some of the plots, particularly non-Aurignacian layer III”, Aurignacian layer IIIb-d
and Middle Paleolithic layers VIII and XI-XIV, slightly resemble the reverse utility curve.
The reverse utility curve reflects an abundance of low-utility elements and
underrepresentation of high-utility elements, suggesting the preferential transport of lowutility elements, possibly as a result of opportunistic scavenging by hominins. It must be
noted, however, that Middle Paleolithic layers VIII and XI-XIV have evidence of densitymediated attrition so it is possible that post-depositional bone loss accounts for the bias in the
249
Table 6.13 Spearman’s rank-order correlation values for fallow deer %MAU and caribou utility
indices (from Binford 1978). *Statistically significant relationships.
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Figure 6.7 Scatter plots of %MGUI and %MAU values for different layers at Klissoura Cave 1
(following Binford 1978).
251
representation of remains. The Upper Paleolithic layers (III” and IIIb-d) may simply
resemble the reverse utility curve because of low incidences of two high-utility elements (the
two points in the bottom right of each plot), as no other lines of evidence indicate that lowutility elements were preferentially transported during these two periods (see below).
Broughton (1999) introduced the mean food utility index (FUI) as a method for
assessing the transport of high-utility body parts to an archaeological site. He uses FUI
values derived by Metcalfe and Jones (1988) for caribou, which are based on Binford’s
(1978) data. FUI is equivalent to MGUI and for this reason is closely correlated with the
meat index (see above). Following Broughton (1999), mean FUI values are derived for
fallow deer in each layer at Klissoura Cave 1 (Appendix J). This is done by multiplying
fallow deer MNE for each element by the element’s respective FUI value (from Broughton
1999). Values for all elements in a layer are then summed, and divided by the total MNE for
the layer to determine mean FUI. Note that Broughton (1999) multiplies FUI by relative
skeletal abundance (RSA), which is an NISP-based value, and MNE is used in this study.
Mean values are plotted in Figure 6.8; the horizontal line indicates mean FUI for a complete
skeleton (Appendix J). Only one layer has a higher mean FUI value than that for a complete
skeleton, which may indicate increased transport distances and the preferential movement of
high-utility elements. Based on mean FUI, during most periods, high-utility elements were
not preferentially brought to the site, suggesting that fallow deer may have been hunted near
Klissoura Cave 1, though density-mediated attrition may be a factor in some layers (see
Chapter 4). Another issue is that transport goals may not have been based strictly around
meat utility, which is explored further below.
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Figure 6.8 Mean FUI for each layer, following Broughton (1999). Horizontal line indicates the
average FUI for a complete cervid. Data from Appendix J.
Body Part Profiles
A final method for interpreting prey transport decisions is the analysis of body part
representation at an archaeological site. Following Stiner (1991), the prey body plan is
collapsed into anatomical regions that are logical packages for the purpose of meat transport
and discard, or dismemberment. In the case of cervids or bovids, nine regions are considered:
antler or horn, head, neck, axial skeleton, upper front limb, lower front limb, upper hind limb,
lower hind limb and feet (see Appendix K for elements included in each region). The hare
body plan includes eight regions, the ones mentioned previously, excluding antler/horn
(Appendix L; Munro 2001). Partridge bodies are divided into eight regions: head, neck, axial,
pectoral girdle, wing, upper hind limb, lower hind limb and feet (Appendix M; Munro 2001).
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Minimal animal units (MAU) (Binford 1978) or standardized MNE (Stiner 1991), which are
equivalent units, are determined by dividing the observed MNE for an element or anatomical
region (regardless of body size) by the expected MNE value for that same element (or
region) in a complete skeleton. This is done for different ungulate species, hare and
partridges in each of the stratigraphic layers with sufficiently large sample sizes. When the
results are plotted as bar charts, a complete anatomical representation is represented by bars
of equal height for all body regions. Dividing the body in this way also bypasses many of the
problems associated with variable structural density because each region contains
corresponding high and low-density portions (Stiner 2002:982; 2004:129; 2005:182).
Similarly to what was done in the taphonomic analysis in Chapter 4, hare remains are
combined with all small mammal axial element data, and all medium ungulate axial elements
are combined with fallow deer data. This done because hare is by far the most common small
mammal and fallow deer dominates most of the assemblages; it is likely that less diagnostic
axial elements came from these species. In the Aurignacian layer IV, where ibex are also
well-represented, medium ungulate axial elements are split between fallow deer and ibex
proportional to the representation of each species in the layer by MNE. In this layer red deer
data are combined with large ungulate axial elements (Appendix N).
Small Game Body Part Profiles
Body part profiles for partridge from Mediterranean backed bladelet industry layer
III’ and the upper Aurignacian layer IIIb-d are presented in Figure 6.9 (data from Appendix
M). Partridge remains in both layers are dominated by elements from the pectoral girdle and
lower hind limb elements; feet, heads and necks are absent in the assemblage. This is similar
254
to what is reported by Bocheński and Tomek (2010); they find high frequencies of partridge
tibiotarsi, humeri, coracoids, pelves, ulnas and tarsometatarsus by NISP. Their study included
a larger sample of avian remains from Klissoura Cave 1 than the assemblage presented here,
though the sample included in this study is representative of all layers from the site.
Figure 6.9 Anatomical profiles for partridges in the Mediterranean backed-bladelet industry (III’) and
upper Aurignacian (IIIb-d) (data from Appendix M).
Hare body part profiles are presented in Figure 6.10 (data from Appendix L) for most
Upper Paleolithic and later layers (excluding the non-Aurignacian Upper Paleolithic layer
III” and the Early Upper Paleolithic layer V) and Middle Paleolithic layer XI-XIV. Recall
from Chapter 4 that density-mediated attrition on hare remains was only a potential factor in
the Mesolithic (3-5a) and Epigravettian (IIa-d) layers (Table 4.1). Indeed, a Kolmogorov-
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Smirnov (K-S) test on MAU in Mesolithic layer 3-5a indicates that hare anatomical
representation is not uniform (Table 6.14). This is also the case for Aurignacian layer IV.
Even in layers where the data do not differ from a uniform representation, hare
Figure 6.10 Anatomical profiles for hares in the Upper Paleolithic and later layers and Middle
Paleolithic (XI-XIV). Note the lack of foot elements in the UP and Mesolithic and neck and axial
elements in all layers (data from Appendix L).
Table 6.14 K-S tests for hare body part profiles by layer. *Asymp. Sig. > 0.05, data do not differ
from a uniform distribution.
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neck, axial and foot elements are poorly represented, particularly in Upper Paleolithic and
later layers. The representation is slightly less biased in the Middle Paleolithic (XI-XIV),
though this layer exhibits an overrepresentation of upper hind limbs and a lack of axial
elements. This observation may contradict the assumption that hare carcasses were brought
to the site complete. Alternatively this pattern may be due to limited spatial sampling in the
excavations (see Chapter 7).
Figures 6.11 and 6.12 provide a more detailed element-by-element comparison of
foot and neck/axial bone representation for hare from Klissoura Cave 1. In both figures,
elements are plotted in descending order of the most dense portion of each element; these
figures illustrate the range of structural densities found in these portions of the hare skeleton
(see values in Pavao and Stahl 1999; Appendix A). There is no bias in the representation of
foot elements with respect to structural density (Figure 6.11) in the assemblages, with the
possible exception of the Mesolithic (3-5a). In general, the entire foot region of hares is
almost entirely absent throughout the later part of the sequence. Hare foot elements are not as
severely underrepresented in the Middle Paleolithic (XI-XIV) (Figure 6.11). Likewise, the
lack of neck and axial elements in the Upper Paleolithic and later assemblages cannot be
explained by variation in bone density (Figure 6.12). Examining the post-cranial axial
skeleton indicates that the innominate is consistently more common than other elements in
the Upper Paleolithic and later layers, though it is not the densest element. Axial elements
from the Middle Paleolithic (XI-XIV) are extremely scarce. Crania and mandibles are
included in Figure 6.12 because they are not as severely underrepresented as the postcranial
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axial skeleton in most layers (Figure 6.10), even though they could be removed during
skinning or transport.
Anatomical profiles for hares are surprisingly uneven, with feet and postcranial axial
elements generally missing from the assemblages. It was established in Chapter 4 that
density-mediated processes cannot explain the lack of hare elements in all Upper Paleolithic
and later layers at Klissoura Cave 1, and discussions above indicate that foot and vertebral
elements with the highest structural density values are no more common than those with low
structural densities. Either hare carcasses were not transported to the site in a whole state or
the excavation did not include all processing and refuse areas. Because hare foot bones are
small, it also is possible that they were not retrieved during excavation or screening. This
may be the case for phalanges, but probably does not explain the lack of metapodials, which
are comparatively quite large.
Cochard and Brugal (2004) argue that lagomorph feet may be absent from
archaeological assemblages because they were removed at the kill site, or because they were
deposited in an area of the site not used for cooking. The presence of the innominate and
head parts, along with an absence of vertebral and rib elements in the Klissoura Cave 1
assemblages may be the result of postcranial axial elements breaking down into
unidentifiable fragments (Cochard and Brugal 2004), whereas innominate fragments
remained diagnostic. In fact, the general composition of hare body parts in the Upper
Paleolithic through Mesolithic is similar to the representation found in the “zone de
préparation culinaire” described by Cochard and Brugal (2004), with the exception of
abundant long bone epiphyses at Klissoura Cave 1.
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Figure 6.11 Plot of the foot region of hares from Klissoura cave 1 by layer. MAU (or standardized
MNE) values for elements included in the foot are presented in order of descending structural density
from the most dense scan site of each element. The expected MAU is presented based on the highest
MNE value for all skeletal elements for each layer. Bone density values from Pavao and Stahl (1999).
Figure 6.12 Plot of the axial region of hares from Klissoura cave 1 by layer. MAU (or standardized
MNE) values for axial elements are presented in order of descending structural density from the most
dense scan site of each element. The expected MAU is presented based on the highest MNE value for
all skeletal elements for each layer. The head region is also plotted to indicate that in some layers it is
present though other axial elements are not. Bone density values from Pavao and Stahl (1999).
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Partridge body part profiles are even more biased than hare remains, but the sample
sizes are large enough to analyze only two layers. It is difficult to determine if this is related
to field butchery, spatial variation in site use, or taphonomic processes.
Ungulate Body Part Profiles
Preferential transport of ungulate elements from anatomical regions based on
nutritional return, or food utility, is of central importance to this discussion. In order to
quantify food utility, I turn to GUI values provided by Binford (1978). Values used here were
originally calculated for caribou, which, as a cervid is the closest species to fallow deer for
which utility values are derived. GUI is preferred over MGUI in this study because
preferential transport is the main question. GUI correlates significantly with the meat index
(see above), though it also takes into account marrow and bone grease, so inherent in using
GUI is the assumption that meat was the main goal of foraging excursions, which may not
always have been the case. For each of the nine ungulate anatomical regions, GUI values are
summed for elements in a given anatomical region (Appendix H). Ungulate anatomical
regions are arranged in order of increasing GUI values. Note that previous presentations of
portions of these data (Starkovich 2009; Starkovich and Stiner 2010) did not arrange
anatomical units in this order.
Fallow deer body part profiles, which includes data for indeterminate medium
ungulate axial elements, are presented by layer in Figure 6.13 in ascending order of GUI
values (from Binford 1978; data from Appendix H). K-S tests indicate that fallow deer
anatomical representation is uniform in layers Mediterranean backed bladelet layer III’,
Aurignacian layer IIIb-d, and Middle Paleolithic layers VIII and XXa-XXb (Table 6.15). Not
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surprisingly, antlers, which have the lowest cumulative GUI values of any anatomical region
are poorly represented in all layers. This may also be due to hunters targeting fewer adult
males, or hunting during the summer, after antlers are shed. The upper hind limb region,
which has the highest cumulative GUI value of any anatomical region, is well-represented
only in the Mediterranean backed bladelet industry layer (III’), the middle Aurignacian (IIIeg) and Middle Paleolithic layer XVIII-XIX (Figure 6.13). These layers also contain high
numbers of lower hind limbs, which may have been attached to the upper limbs during
transport. Three assemblages, from the lower Aurignacian (IV), Middle Paleolithic layer VIII
and Middle Paleolithic layer XXa-XXb, are unusual because they contain high numbers of
lower front limbs, without corresponding frequencies of upper front limbs (Figure 6.13).
Density-mediated attrition may be a biasing factor in the Middle Paleolithic layers, but a
similar relationship does not exist for Aurignacian layer IV (Chapter 4). The lower
Aurignacian is unusual in other ways, however: tooth to bone-based MNE counts indicate an
extreme bias toward teeth in this layer (Chapter 4), and it has the widest range of ungulate
species of all sizes (Chapter 5). Figure 6.14 depicts the anatomical representation of ibex and
red deer in the Aurignacian layer IV (data from Appendix N). Both ibex and red deer have a
more uniform representation of anatomical regions as compared to fallow deer in layer IV
(Table 6.15). Horn and antler elements are lacking from both taxa, as are axial elements for
red deer and upper front limbs for ibex. Lower utility regions tend to be more heavily
represented for both taxa (Figure 6.14).
GUI values for the nine ungulate anatomical regions fall into four categories: low,
medium, high, and highest utility values (Table 6.16). Because the differences among these
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Figure 6.13 Body part profiles for fallow deer in the layers with large sample sizes, ranked by
increasing GUI values (data from Appendix K).
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Table 6.15 K-S tests for ungulate body part profiles by layer. *Asymp. Sig. > 0.05, data do not differ
from a uniform distribution.
Figure 6.14 Body part profiles for red deer and ibex in lower Aurignacian layer IV, ranked by
increasing GUI values (data from Appendix N).
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four categories are quite large, a second set of body part profiles were constructed based on
aggregating the nine anatomical regions into four groups of increasing utility. This provides a
clearer method for determining if higher or lower utility regions were transported to
Klissoura Cave 1. If foragers selectively transported body parts based on their general utility,
a greater representation of high-utility anatomical regions is expected in the archeofaunas.
Figure 6.15 presents medium ungulate body part profiles aggregated by cumulative
GUI values (from Binford 1978). It is striking that the highest utility regions based on GUI,
specifically the upper hind limbs and axial skeleton, are poorly represented in all layers. This
is not surprising, given the lack of axial elements in the assemblages (Figure 6.13), and may
be indicative of post-depositional attrition. Alternatively this could reflect processing
strategies at the kill; perhaps meat was removed from the axial skeleton and transported to
the site, while bulky axial elements were abandoned in the field. The aggregated body part
profiles for most layers are dominated by high-utility regions, followed by medium- and lowutility regions. Medium utility anatomical units are dominant in Aurignacian layers IV and
Table 6.16 Cervid anatomical regions (following Stiner 1991), ranked in order of GUI values derived
for caribou (from Binford 1978).
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Figure 6.15 Body part profiles for fallow deer in the layers with large sample sizes, collapsed into
regions with low, medium, high and highest GUI values (from Table 6.16).
Middle Paleolithic XV-XVII. This may reflect the importance of an additional resource: bone
marrow. The aggregated body part profiles discussed here are based on the GUI, which
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correlates closely with the meat index (Table 6.12). Marrow index values, on the other hand,
are highest for the lower front and lower hind regions (Appendix H), which were aggregated
into the GUI-based medium- and high-utility regions, respectively. It seems, therefore, that
body part transport decisions may have been based on more than just quantity of meat, at
least during some time periods.
Patterns of ungulate carcass transport and butchery differ markedly from those for
smaller game animals. Whereas small game animals can easily be transported to a site in
whole form, the transport of ungulate body parts often has several motivating factors
including the value of meat, bone marrow and bone grease. In zooarchaeology, the specific
skeletal elements are by necessity used as a proxies for the presence of meat. However, field
butchery strategies may involve removing portions of meat from bulky skeletal elements that
are never introduced to a habitation site. Further, the desire for certain nutritional
requirements (e.g., fatty bone marrow during lean times) or availability of specific
technologies (e.g., vessels for bone grease rendering) may expand the goals of foraging
excursions beyond simple meat procurement. There is no clear preference at Klissoura Cave
1 for the transport of the highest-utility elements, though high- and medium-utility elements
(based on GUI) are consistently present in the site during all layers.
MORTALITY PROFILES
Establishing ungulate mortality profiles in an archaeological assemblage provides
information related to herd structure and human hunting strategies, as well as the seasonality
of the occupation of a site. Several lines of evidence yield complementary data, including
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tooth eruption and wear patterns, long bone epiphyseal fusion, and the presence of fetal or
neonate remains in an assemblage.
Age profiles are constructed using eruption and wear schedules for teeth, which are
well-established for many mammals (Deniz and Payne 1982; Grant 1982; Hillson 2005;
Lowe 1967; Magnell 2006; Severinghaus 1949). The dental ages of ungulates are estimated
from the mandibular deciduous fourth premolar (dP4) or the mandibular fourth premolar
(P4). It is possible to use the mandibular third molar (M3) in place of P4, though the use of
the dP4 and P4 is preferred in this study because the deciduous tooth must be shed before the
adult tooth comes into wear, which is not always the case with the M3. Additionally, the dP4
and P4 are highly diagnostic elements even when broken. Tooth side is taken into account,
and only permanent teeth that have some occlusal wear are considered among the permanent
tooth specimens to avoid double counting of individuals (Stiner 1990). Wear stages follow
the three-cohort system presented in Stiner (1990), with the age cohort data collapsed into the
following categories: juvenile, prime-aged adult and old adult. The juvenile stage includes all
animals that died between the time of birth and the shedding of their dP4. The division
between prime-aged adults and old adults is set at approximately 65% of the potential life
span (see Stiner 1990). No attempt is made in this system to assign numeric ages to tooth
specimens. Tooth eruption schedules are not known for all ungulate species at Klissoura
Cave 1, so similar species are used as proxies. Standards for white-tailed deer, published by
Severinghaus (1949), are applied to fallow deer and data on red deer are from Lowe (1967).
Developmental data from Hillson’s (2005) discussion of domestic cattle are applied to
auorchs, data from Angora goats provided by Deniz and Payne (1982) are used for ibex and
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chamois, and data from Magnell (2006) are applied to wild pigs. Tooth eruption and wear
data are available in Appendix O.
Figure 6.16 Tripolar graph indicating living structure and mortally models in an ungulate death
assemblage.
Once ungulate tooth wear stages are established and individuals are organized into the
three-cohort system, the relative proportions of ungulate of different ages are plotted as
percentages of 100 in a tripolar graph. This approach allows comparison of the
archaeological data to known living structures and mortality patterns (Figure 6.16, following
Stiner 1994; 2005). Generally, hominin hunters tend to target prime-aged adult ungulate prey
or to choose age groups randomly upon encounter. This pattern was well-established early in
the Middle Paleolithic and spans into the Holocene in many regions (Adler and Bar-Oz 2009;
Adler, et al. 2006; Enloe 1997; Gaudzinski and Roebroeks 2000; Hoffecker, et al. 1991;
Klein and Cruz-Uribe 1996; Pike-Tay, et al. 1999; Speth and Tchernov 1998; Stiner 1990,
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1994, 2005). In certain situations, particularly during the Natufian in the Levant, human
hunters relied more heavily on juvenile animals, which in some instances is linked to human
hunting pressures and resource stress (Davis 1983; Munro 2001, 2004; Speth and Clark
2006), although seasonality and other motives (such as skin-hunting, see Stiner 1990) must
also be taken into account.
Tooth-based age data for ungulate species are presented in Appendix O; Table 6.17
contains an abridged version of these data. Sample sizes by layer are too small to reach any
firm conclusions regarding age structure at Klissoura Cave 1 by cultural layer, but collapsing
data from the Middle Paleolithic layers and the Upper Paleolithic (and one Epigravettian
specimen) into two groups indicates somewhat different hunting patterns between the two
periods. Relative proportions of juvenile, prime-age adult and old adults exploited in the two
periods at Klissoura Cave 1 are plotted on tripolar graphs in Figures 6.17 and 6.18. Figure
6.17 includes all ungulate species at the site and Figure 6.18 only includes fallow deer.
Ninety-five percent confidence intervals that take into account sample size are included in
the plots (following Weaver, et al. 2011). Two plots are presented because the living
structure of fallow deer differs from many other ungulate species (e.g., Stiner 2005:Fig.
11.11). In both cases, prime-aged adult animals were targeted ubiquitously in the Middle
Paleolithic layers, while the Upper Paleolithic and later layers have a non-selective mortality
pattern. There is little overlap in the confidence intervals of the samples, in either plot,
indicating different hunting strategies in the two periods.
Another method for determining age-at-death is examining fusion states of long bone
epiphyses. Long bone elements fuse in a known, predictable sequence for many species.
Fusion data are useful in understanding juvenile age structures at Klissoura Cave 1 and
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Table 6.17 Age distribution for ungulates in the Klissoura Cave 1 assemblages, based on tooth
eruption data (see Appendix O for more detail).
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Figure 6.17 Proportions of ungulate age groups exploited at Klissoura Cave 1 during the Middle and
Upper Paleolithic based on tooth eruption and wear. During the Middle Paleolithic, prime-aged adult
animals were targeted. More juveniles were incorporated in Upper Paleolithic subsistence pursuits.
Circles indicate 95% confidence intervals (following Weaver et al. 2011) (data from Table 6.17).
Figure 6.18 Proportions of fallow deer age groups exploited at Klissoura Cave 1 during the Middle
and Upper Paleolithic based on tooth eruption and wear. During the Middle Paleolithic, prime-aged
adult animals were targeted. Upper Paleolithic patterns are closer to the living structure of fallow
deer. Circles indicate 95% confidence intervals (following Weaver et al. 2011) (data from Table
6.17).
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helping to establish the season during which a site was occupied. Though all available fusion
data for ungulates are presented in Appendix P, five elements are the focus of the analysis:
proximal first phalanx, distal tibia, distal calcaneum, proximal femur, and distal metapodials
(following Stiner 2005:211, though radii are excluded due to small samples in Upper
Paleolithic layers). Percentages of unfused specimens are calculated by element and the
values plotted in Figure 6.19. Middle Paleolithic layers and the Upper Paleolithic and later
layers were combined into two groups, respectively, to create more robust samples, though
this may mask variation within each time period.
As with tooth eruption data, fusion schedules are not known for all of the ungulate
species in the Klissoura Cave 1 assemblages, so in some cases related taxa are used as
proxies. Fallow deer fusion data are from Kersten (1987) and the reference data applied to
red deer come from Schmid (1972). Bone fusion data applied to ibex are taken from
Noddle’s (1974) study of domestic goats, fusion data from domesticated equids (from Silver
1969) are used for wild ass, and data from domestic pigs (Schmid 1972; Silver 1969) are
applied to wild pigs.
Table 6.18 lists fusion stages for selected fallow deer long bones (Stiner 2005:210).
As with aging data based on tooth wear and eruption, layer-specific fusion sample sizes are
small so the Middle Paleolithic and the Upper Paleolithic and later layers are again combined
into two groups. These data are then graphed in Figure 6.19, which is comparable to Figure
11.12 in Stiner (2005:211). No major differences are apparent between the Middle and Upper
Paleolithic layers based on the state of fusion of fallow deer specimens.
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Figure 6.19 Percentages of juveniles based on five selected elements, based on epiphyseal fusion,
following Stiner (2005), figure 11.12. Selected elements fuse between 12 and 24 months (Kersten
1987). No major differences exist between the ages of juvenile animals exploited in the Middle and
Upper Paleolithic (data from Table 6.17).
Infant and neonate individuals are also represented in the death assemblage. The
small size of fetal/neonate individuals probably belong to unborn animals, based on
comparisons to specimens in known-age collections (Figure 6.20). Fetal or neonate remains
by layer are combined in Table 6.19; most of the remains are from fallow deer. Proportions
of adult to fetal or neonate fallow deer remains are plotted in Figure 6.21. There seems to be
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Table 6.18 Fusion schedules for five elements for fallow deer. Note radii are excluded from Figure
6.19 because of small samples in the Upper Paleolithic layers.
a higher proportion of adult to fetal or neonate fallow deer in the Middle Paleolithic layers,
though the trend is not quite significant (n = 9, rs = -0.633, p = 0.067).
Several interesting points are apparent concerning mortality profiles at Klissoura
Cave 1. Ungulate mortality profiles are instructive in establishing seasonality of occupation,
as well as shifts in the ages of individuals targeted by human hunters. Antler is present
mainly in the Upper Paleolithic and later layers. Male fallow deer possess antlers from
roughly July to April (Chapman and Chapman 1975), so some individuals in the assemblage
were hunted during these months. However, antler may also have been collected and curated
over long periods for tool manufacture, though the presence of some unshed antler bases still
attached to crania indicates that this was not always the case.
Fetal remains recovered in all layers of the site are extremely small compared to
modern stillborn specimens (Figure 6.20). Fallow deer and ibex typically give birth in May
or June (Braza, et al. 1988; Schaller 1977; Spiess 1979), so the size of the fetal remains
indicates that the site was used during the winter or early spring. Similarly-sized fetal
specimens are found throughout the Middle and Upper Paleolithic layers, so the shelter was
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Figure 6.20 Comparison of fallow deer fetal remains from Klissoura Cave 1 and a modern stillborn
goat. (top) Fetal femur from Upper Paleolithic disturbed zone (6-7a), (bottom) Fetal humeri from
Middle Paleolithic layers (VII) (bottom) and (XVa-XVI) (top).
used during the cold months throughout its occupation history, though it could have been
occupied at other times as well.
Ungulate mortality data based on tooth eruption and wear indicate generally different
exploitation patterns between the Middle and Upper Paleolithic layers, with a dominance of
prime-aged adult ungulates in the Middle Paleolithic, and a non-selective pattern in the later
periods. Both cases fit comfortably within the range of observed ungulate mortality patterns
for the Middle Paleolithic through Epipaleolithic across the Mediterranean, including
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patterns specific to fallow deer (Stiner 2005:206). Examining mortality profiles from sites in
Italy, Israel, Lebanon and Turkey does not indicate a clear shift in ungulate hunting patterns
in later time periods, until the Neolithic (Stiner 2005). However, Klissoura Cave 1 provides
an interesting contrast, as the same ungulate species (fallow deer) dominates the assemblages
throughout most of the sequence. The shift from a prime-dominated to non-selective
Table 6.19 Fetal elements identifiable to species by layer.
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Figure 6.21 Proportion of fetal to adult fallow deer by MNI. Data from Table 6.19.
mortality pattern could represent mild pressure on deer populations in the later periods or it
might relate to how hunters exploited herd structures (e.g., focusing on bachelor males versus
maternal herds).
CONCLUSIONS
Human modifications on the faunal remains from Klissoura Cave 1 are apparent
throughout the sequence. The assemblages were principally collected and altered by human
hunters as they targeted prey, transported carcasses to the site, and butchered the remains.
The zooarchaeological evidence clearly indicates that Klissoura Cave 1 was used largely as a
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base camp or residential site for the duration of its occupation. However, some of the
occupations were more intensive than others (see Chapter 7). Small and large game animals
were treated differently with respect to transport and butchery.
Changes in butchery patterns and transport decisions offer only subtle, if any,
indicators of resource intensification at Klissoura Cave 1 from the Middle Paleolithic through
Mesolithic periods. Much of the variation, such as changes in the frequency of burning on
faunal remains, may be explained by changes in site use, which are addressed in Chapter 7,
or limited horizontal sampling during the excavations. Some shifts in butchery patterns, such
as an increase in transverse fractures on ungulate long bones, point to more systematic
butchery efforts or standardization in later time periods. One pattern that is clearly different
between the earlier and later periods is the selection of individuals by age within a herd.
Based on central-place foraging models, it is expected that foragers utilizing
Klissoura Cave 1 would transport higher-utility elements to the site, the further away they
traveled to procure game. According to several analyses presented above, foragers did not
preferentially transport the highest-utility elements during any time period. This suggests that
either foragers at the site did most of their hunting nearby, or that the GUI or FUI are not the
best proxies for ranking elements based on their utility. For example, axial elements have
some of the highest utility values in an ungulate skeleton (Appendix H; Binford 1978).
However, the axial skeleton is rather bulky and may be abandoned in-field. Vertebrae are
useful for bone grease rendering, but might not be transported in the absence of this
technology. There is no evidence for bone grease rendering at Klissoura Cave 1. Further,
nutrient-rich bone marrow may be an important goal of a foraging excursion, and bone
marrow was routinely processed at Klissoura Cave 1 in all time periods. This is particularly
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the case in Aurignacian layer IV, when ungulate third phalanges were routinely opened for
marrow, despite their low nutritional yield. Interestingly, this is one of the layers at the site
with a positive correlation between %MAU and the bone marrow index.
Following the patch-choice model, when Aurignacian layer IV was deposited, the
marginal benefit from intensively processing bone marrow provided a higher return than the
combined travel and handling costs of pursuing the next kill. This may indicate increased
travel distances at this time, though this is not evidenced from body part transport strategies
(though marrow-rich elements were preferentially transported during this period). The patchchoice model may not be relevant to the absence of bone grease rendering at Klissoura Cave
1, though the exploitation of low-utility bone grease often indicates resource stress or
intensification. Resource stress is indicated in certain periods based on intensive marrow
processing (e.g., Aurignacian layer IV) or the exploitation of low-return species (e.g., the
Mesolithic and Epigravettian). However, no evidence has been found for bone grease
rendering as early as the Aurignacian, and Mesolithic and Epigravettian occupations were too
ephemeral at Klissoura Cave 1 to test for grease rendering.
Middle Paleolithic hunters at Klissoura Cave 1 targeted the same ungulate species as
were exploited during later periods, but they chose a different subset of the population. More
fetal or neonate remains were identified in the Middle Paleolithic layers, so overall pregnant
females were hunted more frequently, before that year’s offspring could be born (but there is
only a weak temporal trend). It is possible that this was the result of occupying the site during
the winter, when females were in better physical condition than males. Season of occupation,
in addition to a focus on maternal herds, may explain the prime-aged bias in the Middle
Paleolithic sample. It is likely, based on the size of fetal remains in all layers of the sequence,
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that the site was utilized during the winter or early spring throughout the sequence. In the
Upper Paleolithic, human hunters exploited ungulates less selectively with respect to age and
possibly with a slight bias to juveniles. Coupled with the inclusion of lower-ranked prey
species in later periods discussed in Chapter 5, this latter observation might provide an
argument for resource intensification linked to the growth of human populations in southern
Greece by the end of the Paleolithic, with important variation within time periods in the
sequence that relates to site use.
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CHAPTER 7: SPATIAL VARIATION WITHIN CULTURAL LAYERS
INTRODUCTION
Spatial variation is often apparent across an archaeological site, reflecting different
activity areas or preservation biases in different parts of the site. Though specific activity
areas are more easily discerned at later archaeological sites with permanent architecture or
more formal structures, lateral variation is documented at some Paleolithic sites (Enloe 1991;
Enloe and David 1989, 1992). At the Magdalenian site of Pincevent in the Paris Basin, for
example, Enloe and colleagues (1991; Enloe and David 1989, 1992) found through refitting
faunal elements that reindeer carcasses were shared across the site, with certain high or lowranking carcass parts tending to cluster around different hearths. This is interpreted as
evidence of provisioning some members of the group who were perhaps unable to hunt on
their own, such as widows or grandparents (Enloe 2003b). Spatial distributions indicative of
sharing are also found at other forager sites, in ethnographic and archaeological contexts (e.g.
Binford 1984a; Waguespack 2001; Yellen 1977).
Admittedly, these cases are single-occupation or short-term sites with extraordinary
preservation. It is unrealistic to assume that a similar degree of resolution will be observable
at a deeply stratified site that represents a palimpsest of behavioral events. However,
examining variation within the archaeological layers potentially serves other functions,
beyond the unique circumstances mentioned above. First, analyzing the spatial distribution of
archaeofaunas at a site is a means by which taphonomic processes (Chapter 4) can be better
understood. An example of this is the identification of a bone-poor zone at Hayonim Cave,
where the faunal remains underwent chemical diagenesis in one area of the site and through
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much of the depth of the sequence (Stiner, et al. 2001). This area likely would not have been
obvious if faunal remains were considered in bulk units by layer and could even have been
misidentified as a cleared activity area. A second function that an intra-site spatial analysis
serves is to evaluate whether different areas of the site tended to be used for specific
activities based on the physical characteristics of the location, such as the preferential
placement of hearths to protect them from wind. In this case we expect to see hearths or other
features constructed in the same area through the years. It is possible that faunal remains are
also unevenly distributed, reflecting distinct activities within a site.
In this chapter, the lateral distribution of faunal remains at Klissoura Cave 1 is
examined in order to determine if there are any areas lacking bone that may be explained by
chemical diagenesis or exposure to weathering (such as outside the drip line of the shelter, or
in other unanticipated bone-poor zones). Variation in site use is also discussed in terms of
hearth building, and whether animal processing was focused in certain parts of the site. Some
archaeological layers, specifically the lower Aurignacian (IV) contain many well defined
features such as clay-lined hearths and a possible stone-lined structure (Karkanas, et al. 2004;
Koumouzelis, Ginter, et al. 2001; Pawlikowski, et al. 2000; Stiner 2010). The use of space in
this layer is examined in terms of these features. Differences in sample size based on
discontinuous archaeological layers or the studied sample are taken into account.
METHODS
Prior chapters have focused on behavioral changes that occurred diachronically
through the Klissoura Cave 1 sequence. This chapter focuses on spatial variation of faunal
materials and features within each layer. The goals of this chapter are to consider horizontal
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variation of taphonomic processes within each cultural unit, and to distinguish behavioral
differences related to site use or site topography. This is done by constructing a series of plan
maps for each archaeological layer that depict a specific set of variables. Each map
represents a different cultural layer (i.e., Mesolithic layer 3-5a) and is based on the 1x1 meter
grid system used to excavate the site (see Chapter 2). The first map of each layer provides a
baseline for understanding the subsequent distribution maps. NISP counts for each 1x1 meter
unit are divided by the total NISP count for a given cultural layer in order to calculate the
percentage of the assemblage that came from each unit. In some cases, the percent NISP
count for an excavated unit is zero, which either indicates that the unit was excluded from
this sample because it was included in an earlier study (Koumouzelis, Ginter, et al. 2001;
Koumouzelis, Kozlowski, et al. 2001; Tomek and Bocheński 2002), or that the layer was thin
or discontinuous in that part of the site. Values for subsequent maps (e.g., green bone breaks
or burning frequencies) are calculated for each 1x1 meter unit. For example, the number of
burned bones from square AA1 from Mesolithic layer 3-5a is divided by the total NISP count
for that specific provenience in order to calculate the percent of burned bones in that unit.
The second map in each set examines the spatial representation of green bone
fractures (spits, transverse, or spiral fractures) across the site to determine if processing
activity areas are apparent, based on the percentage of fractured medium ungulate specimens
in each 1x1 meter square per layer. Medium ungulates were chosen for this analysis because
fallow deer are the dominant large prey animal in Klissoura Cave 1 (Chapter 5). Green bone
fractures are examined as opposed to tool marks because most of the Upper Paleolithic faunal
remains are covered with concretions that obscure the surfaces of the specimens (Chapter 4).
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Fresh, or green bone, fractures may occur during marrow processing or when a carcass is
being partitioned during butchery.
A third plan map for each layer indicates the frequency of burning by 1x1 meter unit
in a given layer. Areas with high frequencies of burning may indicate clusters of hearths or a
discreet dumping area. Conversely, a moderate frequency of burning that is consistent across
the site may indicate a high degree of trampling or movement of artifacts across the site. Plan
maps drawn during the excavation by K. Sobczyk were examined to determine hearth
locations; these are indicated on the burning frequency maps. An “H” appears in the lower
left corner of excavation units that contain hearths in each layer. The number of hearths is not
specified in each unit because hearth features (particularly during Aurignacian layer IV) are
often superimposed upon one another and are difficult to distinguish. No hearth information
was available for Middle Paleolithic layers below VIII. The data from each set of maps
described above are discussed by archaeological layer at Klissoura Cave 1. Note that bones
from squares in column B were not included in any of the faunal samples from this study,
though plan maps indicate that they were excavated in the upper layers (K. Sobczyk, plan
maps); these units may represent the original test excavation.
In the lower Aurignacian (layer IV) a rock-lined structure was identified in squares
AA2, AA3, BB2 and BB3, from 145 to 170 cm below the surface (Koumouzelis, Ginter, et
al. 2001; Stiner 2010). Stiner (2010:298) notes that the majority of shell ornaments from this
layer cluster within the man-made structure. In this layer, burning frequencies and butchery
damage (also based on green bone fractures described above) were compared between the
structure and surrounding areas.
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Some of the most interesting features at Klissoura Cave 1 are the dozens of clay-lined
hearths in the Aurignacian and overlying Upper Paleolithic industries (III’ and III”)
(Karkanas, et al. 2004; Kot 2009; Pawlikowski, et al. 2000). The clay-line hearths co-occur
with simple unlined hearths. Understanding the spatial distribution of artifacts in reference to
these features is a logical step in understanding site use. However, initial comparisons
between faunal remains found in the hearths and specimens found outside the hearths proved
to be inconclusive, simply because very few faunal remains were actually recovered in the
clay-line hearth features. Karkanas et al. (2004) note that fallow deer, hare, great bustard and
rock partridge remains were found associated with the clay-lined hearths. Bird and hare
remains, however, have some of the lowest rates of burning in the entire Klissoura Cave 1
assemblages (see Chapter 6), so it is difficult to argue that these specimens were not
deposited on the features after they were used. In addition, many of the hearths are
superimposed on or overlap one another (Karkanas, et al. 2004; Kot 2009; Pawlikowski, et
al. 2000), so it is difficult to assign faunal materials to one specific hearth. Finally, there is no
single cluster of clay-lined hearths at Klissoura Cave 1 (with the exception of the rock-lined
structure in layer IV, see below) making it difficult to associate presumed activity areas with
parts of the site specifically containing clay-lined features.
RESULTS
The first map of each set is simply an illustration of the excavation units and faunal
sample included in the present study. Squares AA4 and BB4 were excavated in the 2005 and
later seasons, after the author had joined the Klissoura Cave 1 team, so all faunal remains
from these units are included in the analysis. The same goes for the Middle Paleolithic layers,
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which were completely analyzed by the author. Discontinuity of some of the layers is also
apparent, such as the upper Aurignacian (IIIb-d) which does not reach the outer margin of the
site, the Early Upper Paleolithic (Uluzzian layer V) which is has a more obvious contact with
overlying layers outside of the drip line of the shelter, and Middle Paleolithic (layer X) which
is also more obvious toward the drip line (see also Karkanas 2010). The maps show that the
excavation area was reduced downwardly from 14 m2 to an 11 m2 area by Middle Paleolithic
layer XI-XIV, and again to a 9 m2 area in Middle Paleolithic layer XV-XVII. The final two
layers of the excavation only included 6 m2.
Mesolithic layers 3-5a
Results of the Mesolithic spatial distribution of faunal remains are presented in Figure
7.1. The majority of the sample is from the squares at the outer edge of the shelter, at the drip
line and beyond, in three units (Figure 7.1a). Green bone breaks on medium ungulate remains
are considerably more frequent at the eastern end of the excavation, which may indicate that
more processing occurred in this area (Figure 7.1b). This may partially be a product of
sample size, however, since squares AA3 and AA4 only contain 10% of the total sample
from this layer. Green bone fractures may be underrepresented in this part of the site; on the
other hand, square BB4 contains a quarter of the sample and still has comparatively low
frequencies of broken bone. The amount of burning in this upper layer is fairly light and is
highest at the outer margin of the site (Figure 7.1c). Few if any hearths were defined in the
Mesolithic layers, and these are not associated with the squares containing the highest rates
of burning, so any concentrations of mildly burned bone may indicate deflated or eroded
hearths or dumping areas associated with fireplace cleaning.
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Figure 7.1 Plan maps for Mesolithic (3-5a), NISP = 256. (a) NISP distribution, (b) frequencies of
fresh bone breaks, (c) burning. Darker boxes indicate higher frequencies of listed parameters. H =
unit with hearths.
Epigravettian layers IIa-d
The Epigravettian (IIa-d) layers contain the smallest faunal sample in the Klissoura
Cave 1 faunal series, only encompassing three excavation units (Figure 7.2a). All of the
specimens from squares AA4 and BB4 display green bone breaks, but only half of the
remains from AA3 are fractured (Figure 7.2b). Like the situation in the Mesolithic layers, the
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Epigravettian has a low incidence of burning damage (Figure 7.2c) and only a few hearths
were recorded during excavation (K. Sobczyk, plan maps).
Figure 7.2 Plan maps for Epigravettian (IIa-d), NISP = 67. (a) NISP distribution, (b) frequencies of
fresh bone breaks, (c) burning. Darker boxes indicate higher frequencies of listed parameters. H =
unit with hearths.
Mediterranean backed bladelet industry layer III’
The sample from the Mediterranean backed bladelet industry layer III’ is much more
robust than any of the overlying samples, though it is still somewhat limited spatially (Figure
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7.3a). Butchery damage on medium ungulate remains occurs in very high frequencies in all
units (Figure 7.3b). Debris from processing activities was discarded in many parts of the
sampled area, or processing took place in all parts of the site. Green bone breakage has little
relationship with the frequencies of burning damage. Burning frequencies are slightly higher
in this layer and grade from less burned toward the back of the shelter (northwest) to more
burned in the southeastern excavation units, beyond the shelter drip line (Figure 7.3c). Most
of the units included in the faunal sample contain hearth features (K. Sobczyk, plan maps).
Figure 7.3 Plan maps for Mediterranean backed-bladelet industry (III’), NISP = 1,359. (a) NISP
distribution, (b) frequencies of fresh bone breaks, (c) burning. Darker boxes indicate higher
frequencies of listed parameters. H = unit with hearths.
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Upper Paleolithic (non-Aurignacian) industry layer III”
The spatial distribution of faunal remains from the Upper Paleolithic (nonAurignacian) industry in layer III” is presented in Figure 7.4a. The sample is large but
incomplete, as some of the materials were included in earlier studies (Koumouzelis, Ginter,
et al. 2001; Koumouzelis, Kozlowski, et al. 2001). The faunal material included in this study
clusters to the northeastern margin of the excavation. The frequency of green breaks on
medium ungulate remains is high in all areas of the excavation, particularly in the southern
Figure 7.4 Plan maps for Upper Paleolithic (non-Aurignacian) industry (III”), NISP = 1,617. (a)
NISP distribution, (b) frequencies of fresh bone breaks, (c) burning. Darker boxes indicate higher
frequencies of listed parameters. H = unit with hearths.
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and western units (Figure 7.4b). Marrow processing activities may have concentrated in the
southwestern part of the excavation areas, but without full horizontal coverage of the site it is
difficult to be sure. Burning damage is higher in this layer than in the later occupations,
particularly in the outermost excavation units of the cave (AA4 and BB4), in addition to one
of the northeastern units (Figure 7.4c). Burn frequencies are lower in the middle of the
sample area, particularly in unit AA2. Hearths are found in several units included in the study
sample, but do not correspond to squares with high incidences of burning. The frequency of
burning and butchery damage do not relate to one another.
Upper Aurignacian layers IIIb-d
The bulk of the faunal materials from the upper Aurignacian (IIIb-d) come from the
northeastern column of excavation units (CC1 and CC2) (Figure 7.5a). Faunal remains were
absent from squares AA4 and BB3, which may indicate that layer IIIb-d was very thin or
absent in this part of the site. Green bone breaks on medium ungulate bones are very
common in this layer, particularly in the northern part of the excavation (Figure 7.5b). The
latter concentration could indicate a processing area, or butchered bones may have been
discarded in this part of the site. Burning frequencies are low in layer IIIb-d, more along the
lines of layers III’ and above (Figure 7.5c). The row of excavation units toward the back of
the shelter has particularly low incidences of burning. Hearths were found in only one square
in this layer (K. Sobczyk, plan maps).
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Figure 7.5 Plan maps for Aurignacian (IIIb-d), NISP = 1,158. (a) NISP distribution, (b) frequencies
of fresh bone breaks, (c) burning. Darker boxes indicate higher frequencies of listed parameters. H =
unit with hearths.
Middle Aurignacian layers IIIe-g
The spatial distribution of faunal remains from layers IIIe-g is presented in Figure
7.6a. Most of the remains came from two parts of the site, the two squares in row 4 and unit
CC2. Some of the gaps in the faunal map are due to certain materials being excluded because
they were studied previously (Koumouzelis, Ginter, et al. 2001; Koumouzelis, Kozlowski, et
al. 2001). Additionally, one box of fauna was excluded from this study due to time
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constraints during the analysis. Green bone fractures on medium ungulate remains are
common, particularly in the center of the excavated area (Figure 7.6b). Areas with high
incidences of fresh breaks have variable frequencies of burning. The middle Aurignacian
layer exhibits a slightly higher rate of burning than the upper Aurignacian (IIIb-d). Many
hearths were recorded in this layer, across the extent of the site (K. Sobczyk, plan maps).
Burned faunal material is most common in the eastern units that were included in the sample,
as well as in square AA2 (Figure 7.6c).
Figure 7.6 Plan maps for Aurignacian (IIIe-g), NISP = 1,699. (a) NISP distribution, (b) frequencies
of fresh bone breaks, (c) burning. Darker boxes indicate higher frequencies of listed parameters. H =
unit with hearths.
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Lower Aurignacian layer IV
The lower Aurignacian (IV) contains the largest faunal sample of any of the cultural
layers at Klissoura Cave 1, and the material is fairly evenly distributed across the horizontal
extent of the site (Figure 7.7a). A slightly higher concentration of faunal remains is evident in
unit AA2 and in two of the squares directly adjacent to this one. Frequencies of butchery
damage are high in layer IV, particularly in the squares at the margins of the sampled area
(Figure 7.7b). The areas with the highest incidences of green bone fractures on medium
Figure 7.7 Plan maps for Aurignacian (IV), NISP = 3,110. (a) NISP distribution, (b) frequencies of
fresh bone breaks, (c) burning. Darker boxes indicate higher frequencies of listed parameters. H =
unit with hearths.
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ungulate remains are also in the squares with the smallest sample sizes. Burning damage is
also common in layer IV, which is not surprising due to the large number of clay-lined and
simple hearths in the layer. Burned faunal remains are concentrated in square CC1, as well as
in the southeastern units (Figure 7.7c). These are some of the areas with high numbers of
hearths, though hearths are recorded in almost all of the units of the excavation (K. Sobczyk,
plan maps). Some of units with high concentrations of burned bone may represent dumping
areas after fireplace cleaning events.
In addition to the spatial distributions described above, the lower Aurignacian (IV)
also contains a possible rock-lined structure (see Figure 7.8 for a map of the structure drafted
by M. C. Stiner). A set of maps similar to the ones presented for the different cultural layers
were constructed for the arbitrary levels in which the structure was observed (145-170 cm
below the surface) (Figure 7.9; squares containing the structure are highlighted). Faunal
remains tend to concentrate in units outside of the boundaries of the rock-lined structure
(Figure 7.9a). It is evident from the plan maps that the majority of the hearths from these
levels are outside of the squares that include the structure (Figure 7.8; K. Sobczyk, plan
maps). Processing damage in the form of green bone breaks on medium ungulate remains is
somewhat lower in the units that contain the structure (Figure 7.9b), so butchering activities
may have focused outside of this area. There is no distinguishable pattern between the
structure and surrounding units in terms of rates of burning of the faunal remains (Figure
7.9c). This may be because the heavy anthropogenic input of the subsequent Aurignacian
occupations quickly filled the structure area after its use was discontinued.
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Figure 7.8 Drafted plan maps in Aurignacian (IV) from 145 to 180 cm below datum by 5cm cuts.
Hearth features are indicated in black and dark grey, limestone rocks in white. The light grey
background represents sedimentary matrix. The shelter feature is apparent in cuts 145 to 170, where
rocks are common and hearths are absent. From Stiner (2010).
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Figure 7.9 Plan maps for Aurignacian layers (IV) containing the structure, NISP = 2,049. (a) NISP
distribution, (b) frequencies of fresh bone breaks, (c) burning. Darker boxes indicate higher
frequencies of listed parameters. Units containing the structure are outlined in black.
Early Upper Paleolithic or Uluzzian layer V
The Early Upper Paleolithic, or Uluzzian, layer V is thin and discontinuous (Figure
7.10a) (Karkanas 2010). This layer is fairly diffuse toward the back of the shelter, but has a
clean contact with underlying and overlying layers beyond the drip line of the cave
(Karkanas 2010). Not surprisingly, the distribution of faunal remains reflects this (Figure
7.10a). Frequencies of green bone breakage are high across layer V (Figure 7.10b). Butchery
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activity seems to have been higher toward the outside of the shelter, but the limited spatial
extent of the sample could undermine this interpretation. Overall, faunal remains in the layer
display high frequencies of burning, particularly in the northern and western portions of the
sample (Figure 7.10c). These clusters may indicate concentrations of deflated hearths or
hearth cleaning areas, but the sample from excavation unit A3 is small. Hearths were found
Figure 7.10 Plan maps for Early Upper Paleolithic (Uluzzian V), NISP = 222. (a) NISP distribution,
(b) frequencies of fresh bone breaks, (c) burning. Darker boxes indicate higher frequencies of listed
parameters. H = unit with hearths.
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in many units of the Uluzzian layer, but because the layer is so thin, it is unclear if they
actually belong to the hearth-rich lower Aurignacian (IV).
Middle Paleolithic layer VIII
The faunal sample from Middle Paleolithic layer VIII clusters in the western part of
the excavation area, with very few specimens coming from the northern and southern
Figure 7.11 Plan maps for Middle Paleolithic (VIII), NISP = 1,569. (a) NISP distribution, (b)
frequencies of fresh bone breaks, (c) burning. Darker boxes indicate higher frequencies of listed
parameters. H = unit with hearths.
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portions of the site (Figure 7.11a). Incidences of green bone fractures on ungulate remains
are high across the excavation area, with the lowest frequencies in square BB2 (Figure
7.11b). This may indicate that marrow processing and other activities took place across the
site during this period. Hearths were recorded toward the back of the shelter (K. Sobczyk,
plan maps), though none are lined with clay. Rates of burning damage on bone are quite
variable (Figure 7.11c); some squares with high rates of burning contain hearths (i.e., CC1),
while others do not. Perhaps some areas with high frequencies of burning represent areas
where hearth cleanings were dumped. Sample size has little effect on rate of burning.
Middle Paleolithic layer X
Middle Paleolithic layer X has the smallest sample of the Middle Paleolithic layers. It
is a fairly small, discontinuous layer that was only found at the outer edge of the site (Figure
7.12a). Faunal remains from this layer are not very evenly distributed across the site, with a
concentration of materials in unit AA3 (Figure 7.12a). Green bone fracture frequencies on
medium ungulate remains are similar for all units (Figure 7.12b), as are frequencies of
burning (Figure 7.12c). In general, this may be a product of sample size or it may be because
the layer was horizontally mixed.
Middle Paleolithic layers XI-XIV
The faunal remains in Middle Paleolithic layer XI-XIV cluster in excavation unit BB2
and adjacent squares (Figure 7.13a). High rates of medium ungulate bone breakage are
apparent in all parts of the excavation except the middle area (Figure 7.13b), which may
relate to intensive marrow processing in specific areas of the site. The rather uneven
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Figure 7.12 Plan maps for Middle Paleolithic (X), NISP = 241. (a) NISP distribution, (b) frequencies
of fresh bone breaks, (c) burning. Darker boxes indicate higher frequencies of listed parameters.
distribution of bone breakage across the site (particularly as compared to burning
frequencies) indicates that the breakage was probably not from post-depositional processes.
There is a modest rate of burning on the remains across the layer (Figure 7.13c), which may
indicate some amount of horizontal mixing, though fracture patterns suggest otherwise.
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Figure 7.13 Plan maps for Middle Paleolithic (XI-XIV), NISP = 1,779. (a) NISP distribution, (b)
frequencies of fresh bone breaks, (c) burning. Darker boxes indicate higher frequencies of listed
parameters.
Middle Paleolithic layers XV-XVII
Faunal remains in Middle Paleolithic layer XV-XVII seem to follow a northeastsouthwest gradient. The faunal sample is large, and the frequency of remains is higher in the
northern excavation units (Figure 7.14a). Green bone breaks on medium ungulate remains are
nearly the inverse of burning damage, with more intensive processing apparent in excavation
column BB (Figure 7.14b). Frequencies of burning damage are higher in this layer than in
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any other, and the most intensely burned areas are in excavation column A (Figure 7.14c).
This spatial bias probably is not related to any kind of slope wash or sweeping, or else we
would expect similar concentrations for both burning and processing damage. It is possible
that more hearths were located in the southwestern part of the site and butchering activities
occurred outside of these areas, but hearth distribution data were not available.
Figure 7.14 Plan maps for Middle Paleolithic (XV-XVII), NISP = 2,031. (a) NISP distribution, (b)
frequencies of fresh bone breaks, (c) burning. Darker boxes indicate higher frequencies of listed
parameters.
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Middle Paleolithic layers XVIII-XIX
Middle Paleolithic layer XVIII-XIX is anomalous relative to all of the other layers at
Klissoura Cave 1 because it displays even distributions for all parameters examined in this
chapter (Figure 7.15). The sample size for this layer is large, but there is no discernable
concentration of faunal remains (with the exception of perhaps excavation unit AA2, see
Figure 7.15a). Green bone breaks on medium ungulate remains are common in the
assemblage and do not focus in any one part of the excavation (Figure 7.15b). Rates of
Figure 7.15 Plan maps for Middle Paleolithic (XVIII-XIX), NISP = 2,199. (a) NISP distribution, (b)
frequencies of fresh bone breaks, (c) burning. Darker boxes indicate higher frequencies of listed
parameters.
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burning damage are low if compared with the other Middle Paleolithic layers, and no
concentrations of burned materials are evident (Figure 7.15c).
One could argue that the even distribution of faunal remains, burn frequency and
evidence of butchery is explained by the limited spatial extent of layer XVIII-XIX. However,
a similar spatial distribution is not apparent in Middle Paleolithic layer XXa-XXb (see
below), which includes the same excavation units, nor is it apparent in some of the younger
Paleolithic layers that were also limited spatially. I would argue that this evenness represents
a high degree of horizontal mixing in layers XVIII-XIX, possibly from trampling or lateral
sweeping of archaeological materials beyond the units included in this sample.
Middle Paleolithic layer XXa-b
The lowest layer of the Klissoura Cave 1 sequence, Middle Paleolithic layer XXa-b,
is spatially limited but has a reasonable sample size (Figure 7.16a). The faunal remains are
largely concentrated in the northeastern part of the excavation, in units BB1 and BB2 (Figure
7.16a). This spatial bias may indicate that that the sediments dip or slope to the northeast, or
that preservation was a problem in the other units. Green bone breaks on ungulate remains
are high in most of the excavation units, with the exception of square BB1 (Figure 7.16b).
Burning damage is much higher in the southwestern units, in the squares with a lower
representation of faunal remains (Figure 7.16c). It is unclear whether hearths were focused in
this area, as maps from this lowest layer could not be located. Interestingly, the square with
the largest sample (BB1) has the lowest frequency of burning and butchery damage.
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Figure 7.16 Plan maps for Middle Paleolithic (XXa-XXb), NISP = 1,106. (a) NISP distribution, (b)
frequencies of fresh bone breaks, (c) burning, (d) medium ungulate specimens by body region. Darker
boxes indicate higher frequencies of listed parameters.
DISCUSSION AND CONCLUSIONS
The horizontal extent of the Klissoura Cave 1 excavation is quite limited (maximum
17 m2) but in fact the site was not large. Though few spatial biases in material distributions
are apparent, analysis of the horizontal layout of the site by layer is useful on several fronts,
including tests for areas of poor bone preservation and loci of repeated activities due to the
orientation of the shelter. In addition, evaluating the distribution of the faunal sample
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horizontally clarifies the limitations of certain layers. The results allow some modest
conclusions to be drawn.
In terms of site preservation and taphonomic processes, there is no evidence
suggesting that faunal remains in certain parts of the site were less likely to preserve than
those in other areas, though a lack of data on the volume of sediment excavated makes this
difficult to quantify. There is also no indication that remains outside of the drip line of the
shelter, which was probably less protected during certain time periods, were less likely to
preserve. In fact, many layers in these southeastern units contain the bulk of the faunal
materials, sample availability and time limits for primary analysis notwithstanding.
Burning damage on faunal remains across the site indicates no consistent vertical
clustering between layers. Activity locations often changed, possibly each time the shelter
was occupied. The shelter was attractive for habitation, which is clear from the intensity of
use during many periods, as well as the flexible arrangement of features within the site.
In general, a spatial analysis of the faunal remains from Klissoura Cave 1 does not
provide any robust conclusions about site use within the different cultural layers. The results
do confirm, however, the conclusion put forth in Chapter 4 that non-human taphonomic
processes had little if any effects on the assemblage, and those observed were not confined to
specific parts of the site. The spatial analysis also indicates that certain layers, such as Middle
Paleolithic layer XVIII-XIX, have a high degree of horizontal mixing, possibly caused by
trampling or sweeping.
The results of this analysis also add an interesting dimension to our understanding of
hearth features and the Aurignacian (IV) rock-lined structure. To begin with, burning and
butchering frequencies seem to be unrelated to the spatial distribution of hearths in Klissoura
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Cave 1. This is not surprising, since hearths locations are highly variable and often overlap.
Klissoura Cave 1, or most Paleolithic sites for that matter, cannot be thought of as having
long, uninterrupted occupations, even in instances where site use was intense. Each new
occupation would introduce more anthropogenic materials into the site, obscuring previous
features or patterns of use. As for the possible rock-lined structure, butchery damage is less
frequent within the units that contain the structure but burning damage is not, even though
hearths do not overlap with this unit. This indicates that faunal refuse filled the structure after
it was no longer in use.
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CHAPTER 8: KLISSOURA CAVE 1 IN CONTEXT:
LATE PLEISTOCENE SUBSISTENCE CHANGE IN GREECE
AND THE MEDITERRANEAN BASIN
INTRODUCTION
The preceding chapters have presented an in-depth analysis of the zooarchaeological
remains from the Middle Paleolithic through Mesolithic layers at Klissoura Cave 1. There are
several goals for this concluding chapter. One is to determine how the Klissoura Cave 1
faunas fit with our understanding of the artifact assemblages of the site, and changes in site
use throughout the history of its occupation. In this discussion, the uniqueness of the site is
highlighted and placed in the context of the Greek Paleolithic. The second goal is to
determine how subsistence data from Klissoura Cave 1 fit with predictions presented in
Chapter 1 concerning, among other things, resource depression as a response to human
population growth or environmental change. Predictions from prey choice, patch choice and
central place foraging models are considered in particular. Also important is a consideration
of the full suite of human activities represented by the major layers. The final goal of this
chapter is to compare dietary trends at Klissoura Cave 1 during the Late Pleistocene with
faunal assemblages from other sites across the Mediterranean Basin. This last part of the
discussion provides a broader picture of subsistence change and hominin adaptations over
this time span.
KLISSOURA CAVE 1
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Perhaps the most unique feature of Klissoura Cave 1 compared to other Greek
Paleolithic sites is the span of the occupations and intact stratigraphy that preserves robust
Middle and Upper Paleolithic components, as well as thinner Epigravettian and Mesolithic
layers. This allows for large-scale diachronic comparisons of many aspects of site use and
changes in stone tool technologies and subsistence patterns. Klissoura Cave 1 is wellpositioned on the landscape, slightly uphill from a perennial stream at the base of a relatively
narrow gorge, which made the shelter an attractive camp or habitation site. Prey species were
likely targeted as they traveled through the gorge and stopped to drink or feed, or as they
passed through the nearby mouth of the gorge from the Argive Plain. There is evidence that
site use varied through time, with particularly intensive occupations during the Aurignacian.
These and other aspects of site use are expanded upon below.
Only slight changes occurred in the lithic assemblages through the Middle Paleolithic,
with the lowest layers (XVIII, XIX and XX) having a higher proportion of blades and
bladelets than the overlying layers. A few Upper Paleolithic tool types in the youngest
Middle Paleolithic layers that were excluded from this study (VI and VII) (Sitlivy, et al.
2007) likely represent localized mixing with younger layers, which is also apparent from the
presence of small numbers of shell ornaments in these layers (Stiner 2010). No major
changes in faunal use are apparent during the Middle Paleolithic at the site. Hunters focused
primarily on prime-aged adult ungulates, fallow deer in particular, and supplemented their
diets with tortoises and occasionally hares. Human modifications on carcasses are similar
among the Middle Paleolithic assemblages; frequencies of burning and marrow processing
are high throughout. A higher proportion of early-term pregnant female fallow deer were
hunted during the Middle Paleolithic than in later periods, which may suggest that the site
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was occupied primarily during the winter, and that adult females were targeted from maternal
herds because they were in better nutritional condition than males.
The Early Upper Paleolithic or Uluzzian layer V is important for addressing larger
questions concerning the Middle to Upper Paleolithic transition. Layer V represents a rare
occurrence of the Uluzzian outside of Italy and is several thousand years older (>39,000 BP)
than at some of the Italian sites (but see Kuhn, et al. 2010). Unfortunately, the faunal
assemblage from this period is smaller than the other layers (NISP < 300), making many
comparisons difficult. The Early Upper Paleolithic faunal assemblage has some features in
common with the Middle Paleolithic layers, such as comparatively high proportions of
tortoise remains and a dominance of fallow deer, though hares are somewhat more important
in the Uluzzian, similar to the overlying Upper Paleolithic and later layers.
All evidence indicates that Klissoura Cave 1 was occupied intensively during the
Aurignacian period. This is particularly striking since the Aurignacian is absent from most
Greek Paleolithic sites, particularly those in the north. In Peloponnese, Aurignacian deposits
are found at Franchthi Cave and possibly also in Apidima and Kephalari Caves. The
Klissoura Cave 1 Aurignacian lithic assemblage is massive (n > 90,000; Kaczanowska, et al.
2010), and the bulk of the ornaments comes from these layers (Stiner, et al. 2010). The
Aurignacian also contains hundreds of hearth features, dozens of which are clay-lined
structures (Karkanas, et al. 2004; Pawlikowski, et al. 2000), and a possible rock-lined
habitation structure in Aurignacian layer IV (Koumouzelis, Ginter, et al. 2001; Stiner, et al.
2010). With the exception of the faunal remains, artifact frequencies tend to cluster inside of
the structure, while hearths occur only outside of it.
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The clay-lined hearth features are one of the most unique components of Klissoura
Cave 1, but it is difficult to determine their function because they seem to have been cleaned
out after use. Botanical evidence is rare in the features, and the faunal remains found directly
on the hearth surfaces do not seem to be directly associated with them. There is no specific
reason to believe that the clay-lined hearths served a single purpose; they may have been
used for parching plant materials, braising meat, or as brazier-like satellite hearths around a
larger simple hearth. One purpose that they probably were not used for is as shallow vessels
for bone grease rendering, as evidence of this practice is lacking from the site. What is
interesting about these features is their ubiquity throughout the sediments that may span a
few millennia at Klissoura Cave 1. To date they are not found at other contemporary sites in
Greece or the Mediterranean.
The faunal assemblages of the Aurignacian are much more variable than those from
the Middle Paleolithic. Aurignacian layer IV is taxonomically richer than the earlier periods,
probably because of the appearance of mosaic parkland-forest communities in the area
(Albert 2010; Ntinou 2010). Frequencies of hare are also high. In the stratigraphically later
Aurignacian layers, fallow deer are the dominant large game species and low-ranked small,
fast-game were commonly exploited. There is an increase in rock partridge and great bustard
in Aurignacian layers IIIb-d, probably in response to increasingly open vegetation in the
vicinity of the site. Evidence for the exploitation of labor-intensive land snails appears at this
time. Burned bone is common in the Aurignacian layers, and ungulate crania exhibit higher
frequencies of burning than postcrania. Impact and cut-mark data are difficult to quantify in
these layers due to ash concretions on the bone surfaces. Bone marrow processing intensifies
during Aurignacian layer IV, possibly as a result of prolonged site use. An increase in the
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importance of marrow extraction is evidenced by the preferential transport of marrow-rich
elements and more common processing of marrow-poor terminal ungulate phalanges. There
is also a shift in the Aurignacian to a non-selective mortality profile for ungulates, quite
different from the prime-dominated patterns of the Middle Paleolithic. This may be partly
explained by the main season(s) of occupation at the site. Fetal remains in the Upper
Paleolithic layers are similar in size to those found in the Middle Paleolithic, indicating some
winter occupations, but fewer fetal remains in general are found in the Upper Paleolithic.
The cultural affiliations of Upper Paleolithic layers III” and III’ are distinct from the
Aurignacian, but the faunal exploitation patterns of III” do not differ substantially from
variation found in the Aurignacian, with the exception of an increase in land snail
exploitation. In fact the species representation is very similar to that of Aurignacian layer
IIIe-g, with high frequencies of fallow deer and some evidence of hare exploitation, but few
partridges and bustards. A major shift in prey representation occurred in association with the
younger of the two industries (III’), when small fast-moving game overtakes fallow deer by
NISP count for the first time. Great bustard and partridge frequencies increase and slightly
higher rates of European wild ass are found, indicating more open vegetation. Tortoise
disappears from the faunal assemblages after this layer, as does great bustard. Faunal and
lithic densities also decrease at this time relative to the Aurignacian series and layer III”,
possibly indicating a decrease in site use.
Site occupation intensity drops further during the Epigravettian and Mesolithic
periods at Klissoura Cave 1. The lithic assemblages from these periods are modest, and
faunal densities are the lowest of the entire sequence. Sample sizes from the youngest layers
are small, so much so that some of these assemblages had to be excluded from many of the
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analyses. Land snails are uncommon in the Epigravettian layer, but peak in the Mesolithic.
The ungulate component is more even with respect to species representation in the
Epigravettian and Mesolithic layers, because of the decline of fallow deer relative to other
taxa, but the faunas are dominated by hare. Relatively few hearths were excavated and
burned bones are uncommon in these layers.
There is a clear shift in foraging behaviors between the Middle Paleolithic and
Aurignacian layers at Klissoura Cave 1 in terms of prey representation, mortality profiles,
processing intensity, and burning frequencies on certain ungulate body parts (i.e., crania).
Taken in conjunction with the abundance of lithic materials, the large ornament assemblage,
dozens of clay-lined hearths and possible structure, it seems that Aurignacian layer IV
marked the beginning of a period of exceptionally intensive site use at Klissoura Cave 1.
Occupation intensity peaked in layer IV, but remained high throughout the other Aurignacian
units and into layers III” and III’. The second major faunal shift occurred after layer III’,
when tortoises and bustards completely disappear from the assemblage, and hare becomes
the most common prey species at the site. Klissoura Cave 1 was either abandoned during the
Last Glacial Maximum (LGM), or there is a depositional hiatus at this time. Use of the site
during the Epigravettian and Mesolithic was considerably lighter based on all lines of
artifactual and faunal evidence. However, several other caves in the areas were visited by
groups during these periods (Koumouzelis, et al. 2004). Many of the important Greek sites
that date to the Mesolithic are coastal (Jacobsen 1981; Jacobsen and Farrand 1987; Sampson
1998), and evidence for open water seafaring appeared during this period.
Changes in Site Use and Occupation Intensity
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Several different trends are apparent at Klissoura Cave 1 that relate to occupation
intensity and the use of faunal resources. These trends are often at odds with one another and
may reflect a combination of changes in the use of the site itself, as well as larger human
population changes on the landscape of southern Greece. Figure 8.1 illustrates broad changes
in ungulate diversity, the use of small, fast-moving game species, and fluctuations in
occupation intensity through the sequence at Klissoura Cave 1. Diversity data are based on
the reciprocal of Simpson’s Index values presented in Chapter 5, and small, fast-moving
game are from NISP proportions. Changes in occupation intensity are based on faunal and
lithic densities, as well as anthropogenic sedimentation and the relative abundance of hearth
features in different layers. As mentioned in earlier chapters, “occupation intensity” or
“increases in site use” can reflect 1) human groups using a site more often, 2) an increase in
group size at the site, or 3) humans spending more time at an individual site. This study does
not seek to distinguish between these behaviors; an increase in site use may represent one or
a combination of several of these factors.
Ungulate diversity roughly tracks environmental changes, as discussed in Chapter 3;
diversity is generally higher in periods in which climate was wetter (i.e., Aurignacian layer
IV). Diversity values drop when vegetation contracts in dry phases, implying that there are
fewer suitable habitats for ungulate species. It might be expected that the site was used more
often by humans during these phases, when diverse ungulate prey was abundant on the
landscape, but this does not always seem to have been the case. Rather, the intensity of site
use is low through the Middle Paleolithic and peaked during the Aurignacian and later Upper
Paleolithic phases, followed by a decline after the LGM. It is likely that hominin populations
were generally low in the Middle Paleolithic of Greece in general. Greece certainly would
315
Figure 8.1 Illustration of changes through time in ungulate diversity, occupation intensity and small
game use at Klissoura Cave 1.
have been an early area of colonization by modern humans as they moved across Europe,
though the occupation pulse during the Aurignacian in Klissoura Cave 1 post-dates this
phase. It must be noted that no increase in site use is associated with the Early Upper
Paleolithic (Uluzzian) as compared to Middle Paleolithic layers. It is also possible that
southern Greece was a refugium for many plant and animal populations during the mild but
climatically variable MIS 3, making it an attractive region for human habitation, especially
after 40,000 BP. However, no contemporary sites in Greece preserve a similarly intense
Aurignacian occupation, though this may simply reflect the small number of Upper
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Paleolithic sites that have been excavated in the region. The LGM in Greece was either a
period of depopulation or an interval in which sediments did not accumulate (e.g., Farrand
2000; Karkanas 2010). After the deglaciation, Klissoura Cave 1 was occupied much less
intensively during the Epigravettian and Mesolithic periods.
The proportion of small, fast-moving animals in the Klissoura Cave 1 assemblages
does not track the environmental changes or occupation intensity (Figure 8.1). Rather, there
is occasional exploitation of hare in the Middle Paleolithic, although some of these animals
may be intrusive. Small, fast-moving animals (low-ranked prey) become much more
common in the Upper Paleolithic and eventually surpass large game by NISP counts in the
Epigravettian and Mesolithic. It is significant that low-ranked small game was hunted more
and more in later periods, even though occupation intensity had actually declined. This result
likely reflects a regional trend (see also Stiner and Munro 2011), and could indicate that
rising human populations stressed higher-ranked faunal resources (e.g., large game and
small, slow-moving animals) on the landscape in general.
Season of occupation is also relevant in a discussion of site use at Klissoura Cave 1.
Fetal remains throughout the sequence are exceptionally small, indicating that the site was
often utilized during the late winter or early spring in all culture periods. However,
proportions of fetal remains are generally higher in the Middle Paleolithic, as is the
representation of prime-aged adult animals. This pattern may reflect the movement of
hominin populations as part of seasonal rounds and/or the targeted hunting of maternal herds.
Upper Paleolithic and later groups may have occupied the site during a wider range of
seasons, opportunistically hunting maternal and bachelor herds.
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RESULTS FROM THE PERSPECTIVE OF EVOLUTIONARY ECOLOGY
In this section, I return to predictions set forth in Chapter 1 about the faunal record at
Klissoura Cave 1 based on models from evolutionary ecology. One of the most interesting
and valuable aspects of the application of such models is when the archaeological record
deviates from the predictions. Deviations may indicate a unique cultural response or a shortcoming of the model in its initial form.
Prey Choice
Following the prey choice model, it is expected that lower-ranked resources will enter
the diet of foragers when high-ranked resources are less abundant in the environment (Emlen
1966; MacArthur and Pianka 1966; Pianka 2000; Stephens and Krebs 1986). A decrease of
high-ranked resources can be caused by human hunting pressures, environmental change, or
a combination of both. The inclusion of more lower-ranked resources in the diet may
manifest as a generally wider diet breadth, which can result in greater evenness in the
proportions of prey types in an archaeological assemblage. Another marker of expanding diet
breadth is the greater inclusion of low-ranked resources (e.g., small, fast-moving animals)
relative to higher-ranked ones (e.g., small, slow-moving game or ungulates).
At Klissoura Cave 1, a trend in resource depression and widened diet breadth is
apparent from multiple lines of evidence. There is a general increase in prey evenness based
on the reciprocal of Simpson’s Index for all taxa, although the trend is not statistically
significant. The proportion of prey types changes through the sequence, favoring small quick
types and labor-intensive invertebrates with time, thus suggesting an increase in the costs of
capture or processing in relation to food yields. There is a significant increase in the
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proportions of small to large prey animals from the Middle Paleolithic through Mesolithic
periods at Klissoura Cave 1, as well as an increase in the small game fraction of fast-moving
types at the expense of slow-moving species. The decline of high-ranked prey types over
time cannot be explained by the climatic fluctuations, as discussed in Chapter 3. Taken
together, these lines of evidence indicate some degree of resource depletion and widened diet
breadths at Klissoura Cave 1 from the Late Pleistocene through the Early Holocene.
One possible deviation from the prey choice model is the occasional appearance of
low-ranked hares in the Middle Paleolithic, particularly in layers XI-XIV, when there are no
indications of resource stress. However, hare specimens are typically rare in the Middle
Paleolithic assemblages overall (MNI=1 in most layers). Middle Paleolithic hominins were
capable of exploiting hares, but this animal never comprised a significant part of the diet at
Klissoura Cave 1. Layer XI-XIV has a higher incidence of hare remains than any other layer
(MNI=2), but there is reason to believe that at least some of the remains are intrusive. Hare
remains in layer XI-XIV have no evidence of cut marks and frequencies of burning are much
lower than those for the assemblage as a whole. Early exploitation of small, fast-moving
game has been noted elsewhere in the Mediterranean (e.g., Blasco and Fernández Peris 2009;
Sanchis Sera and Fernández Peris 2008), and though hares were occasionally exploited in the
Middle Paleolithic at Klissoura Cave 1, the major trend is still toward the increased reliance
of lower-ranked resources in later periods.
Central Place Foraging
Central place foraging models attempt to explain the transport decisions that foragers
make when bringing food items to a home base (Orians and Pearson 1979; Schoener 1979). It
319
is expected that, if human hunters exploit large game animals near a home base or habitation
site, they will move more than just high-utility portions of carcasses. Rather, both high and
low-utility elements are brought to the site because the cost of transporting the entire carcass
(or large portions of it) is fairly low. The result is a more even anatomical representation at a
site. Conversely, if foragers travel a great distance to hunt, they maximize their returns by
only transporting the highest utility elements back to the habitation site. If local resource
depletion occurs as the result of overhunting or changing environmental conditions, it is
expected that there will be an increasing trend toward high-utility elements in an
archaeological assemblage (Broughton 1999; Nagaoka 2002a; 2005, see also Faith 2007 for
an example of changing transport decisions based on more favorable environmental
conditions). It must be noted, however, that though food utility is based on anatomical
characteristics (i.e., meat or marrow content of different elements), it is a relative concept
that also depends on technologies available to foragers (e.g., bone grease rendering, which
currently does not seem to predate the Gravettian, see Manne and Bicho 2009).
Several different analytical strategies were employed to examine whether there were
shifts in the relative utility of ungulate body parts transported to Klissoura Cave 1, or if body
part transport was more complete in certain time periods. Based on other changes in faunal
exploitation patterns indicative of resource stress (e.g., low-ranked prey exploitation in the
youngest layers or increased marrow processing in Aurignacian layer IV), we might expect
preferential transport of high-utility body parts during certain periods. Evaluating the average
food utility index values for body parts transported to the site in each layer did not yield any
trend that might indicate human hunters exploited large game further from the site in
expected periods. In fact, all layers except for one in the Middle Paleolithic had a lower
320
average FUI value than is found in a complete cervid skeleton. This indicates that there was
no preferential transport of high-utility body parts (based on FUI values) in most periods.
This is also the case when correlations were tested between anatomical representation and
MGUI/GUI values; no relationships indicate that there was an increase in the transport of
high utility elements in later periods. Similarly, examining changes in ungulate body part
representation by anatomical unit does not indicate directional changes in transport strategies
through the sequence, though some layers had more uniform distributions than others based
on K-S tests (e.g., Upper Paleolithic layers III’, IIIb-d, red deer and ibex in layer IV, and
Middle Paleolithic layers VIII and XXa-XXb).
There are several possible explanations for the lack of trend in body part transport
decisions at Klissoura Cave 1. One is a taphonomic bias; density-mediated attrition is a
potential problem, particularly in the Middle Paleolithic layers, which would cause lowdensity, high-utility elements such as vertebrae to be underrepresented in the assemblages.
Another explanation concerns the technologies available to foragers occupying the site.
There is no evidence of bone grease rendering, so it is possible that bulky vertebral elements
were abandoned in the field where the carcass was butchered. Faunal elements at an
archaeological site are used as proxies for meat transport, but in fact edible tissues can be
removed from bones that are then left at kill sites. There may also be an issue with the
assumed goals of foraging excursions, namely that FUI and GUI rely too heavily on the meat
index, when in many cases food transport correlates with marrow-rich elements (see below).
A final explanation for the overall lack of temporal trend in large game transport patterns is
that human hunters at Klissoura Cave 1 simply did not travel greater or smaller distances to
hunt large game across periods. Rather, they may have periodically abandoned the site when
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large game became less abundant. There must be a point where the distance traveled for large
game is so great and transport costs are so high that foragers can instead get a higher return
from exploiting locally available small game (Cannon 2003).
There is evidence for all of these possibilities at Klissoura Cave 1, and to an extent it
may be a combination of factors. However, the position of the site on the landscape, next to a
perennial stream that would have attracted large herbivores, probably allowed foragers to
hunt prey close to the site during most periods. By the end of the sequence, it seems that
human groups employed a different intensification strategy than traveling long distances to
procure game. Low-ranked prey items played a greater role in forager diets during later
periods of the site, and the shelter was certainly used less intensively. By the Mesolithic, it is
also clear that other local sites were in use, including other shelters in the Klissoura Gorge, as
well as coastal sites such as Franchthi Cave, where marine resources were included in the
widening dietary repertoire.
Results on Patch Choice based on Carcass Processing Intensity
The patch choice model predicts which high-ranked patches on the landscape a
forager will exploit (MacArthur and Pianka 1966), and the marginal value theorem
determines how much time foragers will spend in a given patch based on the depletion of the
patch as a function of time (Charnov 1976; Charnov, et al. 1976). One anthropological
application of the patch choice model and the MVT is to butchery or processing intensity of
prey carcasses. In this case resource intensification as reflected in increased bone marrow or
grease processing over time can indicate human population pressure or resource stress
(Burger, et al. 2005; Nagaoka 2005). Changes in processing intensity can be in response to
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seasonal shortages, or if they occur over a period of time may reflect large-scale trends in
carcass exploitation patterns provoked by food shortages.
At Klissoura Cave 1, there is no indication that marrow processing intensified
through the sequence, though there are certain periods where it was somewhat more
important (i.e., Aurignacian layer IV). Ungulate elements with the least amount of marrow
(e.g., phalanges, particularly the third phalanx), show no directional trend in the frequencies
with which they were opened, while other elements were always processed for marrow.
During the formation of Aurignacian layer IV, there is evidence for more intensive
processing of terminal phalanges, as well as a preference for the transport of marrow-rich
elements to the site. This layer also corresponds with an increase in site use, evidenced by
higher quantities of lithics, ornaments and hearth structures. At the same time, ungulate
diversity is high in layer IV, so it seems that human foragers exploited the rich environment
very intensively during this period.
There is no evidence for heat-in-liquid bone grease rendering at the site, which is
considered to be another important measure of intensification (Bar-Oz and Munro 2005;
Manne and Bicho 2009; Munro 2004; Munro and Bar-Oz 2005; Nakazawa, et al. 2009;
Stiner 2003b). Bone grease rendering technologies were probably not part of the cultural
repertoire when the site was occupied most intensively (e.g., the Aurignacian). As of this
writing, there is no evidence for bone grease rendering in Eurasia earlier than the Gravettian
(Manne and Bicho 2009). By the Epigravettian and Mesolithic periods, large game is so rare
at Klissoura Cave 1 that potential changes in processing intensity are nearly irrelevant.
Resource Depression: Climate Change, Human Population Growth or Both?
323
There is some evidence of resource depression in the vicinity of Klissoura Cave 1
beginning in the Upper Paleolithic and increasing in intensity with the Epigravettian and
Mesolithic periods. Very little change in hunting or butchery practices occurred during the
Middle Paleolithic at this site, with hominin groups almost exclusively targeting high-ranked
resources. These resources included tortoises, which have low fertility rates and whose
populations are easily impacted by increased harvesting pressures (Stiner, et al. 2000). By the
Upper Paleolithic and later periods, lower-ranked resources comprised a growing part of the
diet, and the frequency of high-ranked tortoises declined substantially in the Aurignacian.
The rising proportion of prey species that inhabit open areas likely correlates with short-term
fluctuations of vegetation types in the area, whereas other long-term trends do not relate to
environmental conditions. It is interesting, for example, that great bustard was exploited at
low-levels during the Aurignacian but completely disappear from diets by the Epigravettian,
an interval in which open parkland communities were expanding (Albert 2010; Ntinou 2010).
Human hunting pressures may have depressed populations of these slow-reproducing birds.
Tortoises also completely disappear from the diet by the Epigravettian. Because of these
changes, and the fact that ungulate diversity increased again in the Mesolithic while
exploitation of large game species declined, it seems likely that human foragers were a major
cause of resource depletion in the Late Pleistocene and early Holocene in the eastern
Peloponnese.
It is noteworthy that intensified foraging did not extend to increased processing
intensity or longer travel distances for ungulate hunting. Though human population densities
were on the rise in southern Greece, foragers could continue to move along if all local animal
resources became depleted. It could also be that the gorge remained a prime vantage point for
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hunters because of its position at the edge of the Argive Plain. In any case, the relentless
increase in the use of quick small game with time cannot be explained by environmental
changes alone.
KLISSOURA CAVE 1 WITHIN GREECE AND THE MEDITERRANEAN BASIN
At Klissoura Cave 1 there is evidence for resource depression during the Upper
Paleolithic and later periods, particularly when it comes to prey choice. Transport decisions
and processing intensity do not change with time, though there are fluctuations in some
periods. How do the subsistence trends at Klissoura Cave 1 compare with those found at sites
across the Mediterranean Basin? Here I compare general trends in faunal exploitation at
Klissoura Cave 1 to contemporary sites that contain robust data sets from the northern and
eastern Mediterranean. Sites are included based on their location in the Mediterranean zone
defined by Blondel and Aronson (1999:8) (Figure 8.2).
Figure 8.2 Map of the four quadrants of the Mediterranean. Shaded areas indicate Mediterranean
ecosystem. From Blondel and Aronson (1999).
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Faunal data from Late Pleistocene sites in Israel provide a high-resolution picture of
dietary trends in the eastern Mediterranean. In their work on the Kebara Cave Middle
Paleolithic faunal assemblages, Speth and Tchernov (1998; 2001; 2002, see also Davis 1977
for results from earlier excavations that support many points discussed here) conclude that
Neandertal groups utilizing the site were successful hunters of large, prime-aged adult game
animals during the late Pleistocene, including gazelle, fallow deer, red deer, wild pigs and
aurochs. The most common species at the site are gazelle and fallow deer. The authors find
that the hunting of male and female fallow deer and gazelles generally reflects their
proportions in living populations; temporal shifts in their proportions relate to changes in the
season of site occupation (Speth and Tchernov 2001). Neandertals typically transported highutility body parts to the site after low-utility bulky elements were removed and abandoned in
the field. In terms of small game use, Speth and Tchernov (2002) find evidence from burning
patterns that, in addition to their value as a food source, tortoise carapaces may also have
been used as containers. They observe tortoise size diminution at the end of the Middle
Paleolithic, which may indicate mild prey population suppression (e.g., Stiner, et al. 2000),
though the authors note that the early Upper Paleolithic occupations of Kebara Cave were
ephemeral and that other explanations must be explored (Speth and Tchernov 2002).
Expanding on these studies, Speth and Clark (2006) note that frequencies of large ungulates
such as red deer and aurochs declined steadily over time until they comprise a fraction of the
early Upper Paleolithic assemblage. Also, foragers at the site exploited more young fallow
deer and gazelles in later periods (Speth and Clark 2006).
Based on an extensive study of the Hayonim Cave faunal series, and in conjunction
with other pan-Mediterranean work, Stiner (2005) notes three major trends in human
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subsistence strategies over the course of the Middle and Late Pleistocene through Early
Holocene. First is the exploitation of prime-aged adult ungulates that began before the
Middle Paleolithic (Stiner 2005; Stiner, et al. 2009). Next is a general trend in small game
exploitation. Middle Paleolithic hominin groups spent little time pursuing small game
species, except for those with the highest return rates such as tortoises or marine mollusks.
Average tortoise body size at Hayonim and Kebara Caves began to decline after about
44,000 BP, as a result of overharvesting larger, more productive individuals, a trend that
continued through the Natufian (Stiner 2001, 2005; Stiner, et al. 1999; Stiner, et al. 2000).
Low-return small, quick species such as birds become common in the Upper Paleolithic, and
by the Epipaleolithic lagomorphs increased in importance (Stiner 2001, 2005; Stiner, et al.
1999; Stiner, et al. 2000). The final trend is the appearance of more efficient hunting
technologies including stone and bone-tipped projectiles in the late Middle and early Upper
Paleolithic, possibly including snare, net and trap technology late in the Upper Paleolithic
(Stiner 2005). Labor-intensive carcass processing technologies, such as bone grease
rendering, also appear in later periods (Bar-Oz and Munro 2005; Munro 2004; Munro and
Bar-Oz 2005).
By the Epipaleolithic, several lines of evidence indicate that resource intensification
was widespread in the Levant. At multiple sites, evidence for depression of medium ungulate
(i.e., fallow deer) populations began early in the Epipaleolithic and peaked in the Natufian, at
time when small ungulates (i.e., mountain gazelles) overwhelmingly constitute the ungulate
component of the archaeofaunas (Munro 2004, 2009a, 2009b; Stutz, et al. 2009). Though
ungulates remained the dominant prey type in the Levantine sites discussed, the incidence of
small quick game also increased starting in the Early Natufian (Bar-Oz 2004; Munro 2004,
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2009a, 2009b). In addition to changes in the prey species exploited, young gazelles were
commonly hunted during the Early Natufian (Munro 2004, 2009a, 2009b). Bone marrow was
intensively processed across the Epipaleolithic and Natufian periods at the Israeli sites (BarOz and Munro 2007), and there is strong evidence for bone grease rendering at this time
(Munro 2004, 2009a; Munro and Bar-Oz 2005).
In the Mediterranean zone of Turkey, recent work at the Paleolithic sites of Uçağızlı
Caves I and II (Kuhn, et al. 2009; Stiner 2009) and Epipaleolithic Öküzini and Karain Caves
(Atici 2009) provides large faunal data sets from the Late Pleistocene. Uçağızlı Caves I and
II contain complementary Middle Paleolithic (II), Upper Paleolithic (I) and Epipaleolithic (I)
sequences. The most common prey types in the assemblages are fallow deer and bezoar goat,
whose abundances through the sequence oppose one another, depending on climate regime,
followed by roe deer and a range of other ungulate and carnivore species in smaller numbers.
There is a decline in the evenness of ungulate prey through the sequence that follows the
disappearance of large ungulates such as aurochs and red deer (Kuhn, et al. 2009; Stiner
2009). A striking trend is apparent in the small game component for these sites. Small, slowmoving species such as tortoises and shellfish are present throughout the sequence, though
tortoise abundance drops in the Upper Paleolithic. Conversely, small fast-moving
lagomorphs, birds and fish only become important in the Upper Paleolithic and
Epipaleolithic layers (Kuhn, et al. 2009; Stiner 2009). Little variation is noted in body part
transport decisions through the sequence, with axial elements and phalanges consistently
underrepresented. A change in butchery practices occurred during the early Upper
Paleolithic; in earlier periods most cut marks were diagonal to the main axis of elements,
328
whereas in the Ahmarian most cuts have an axial orientation, possibly reflecting the removal
of large, intact portions of meat, possibly for smoking (Stiner 2009).
Epipaleolithic faunal data from southwest Turkey at the sites of Öküzini Cave and
Karain Cave are presented by Atici (2009). The data reveal a nearly exclusive focus on wild
sheep and goat for most of the Epipaleolithic, which Atici (2009) argues to be evidence for
hunting specialization. There is a general diversification of the diet toward the end of the
Epipaleolithic in the area, with the inclusion of low and high-ranked game such as roe deer,
wild boar, tortoises, hares and partridges into the meat diet. The author notes that this dietary
expansion occurred during a time of climatic amelioration and that it may be the result of
increased availability of a wider range of species (Atici 2009).
In Greece, excluding Klissoura Cave 1, the most substantial faunal data sets available
are from Franchthi Cave (Payne 1975; Rose 1995; Stiner and Munro 2011), Kastritsa
(Kotjabopoulou 2001), Klithi (Gamble 1997; 1999), Boïla (Kotjobopoulou 2001) and Cave
of Cyclope (Mylona 2003; Powell 2003; Trantalidou 2003). Of these, Franchthi Cave is the
only site that preserves a long sequence of faunal remains from the Aurignacian through
Mesolithic (and a substantial Neolithic component that is not the focus here). The Franchthi
Cave faunas provide data on long-term subsistence trends. Ungulates such as red deer and
equids dominate the Upper Paleolithic layers of Franchthi Cave, and shifts in their
frequencies track environmental change (Payne 1975; Stiner and Munro 2011). By the
Epigravettian, ungulate diversity declines and the meat diet widens to include small game
animals such as land snails, shellfish, coastal fish species, and pond turtles (as opposed to
tortoises, which are found earlier) (Stiner and Munro 2011). Across the Mesolithic, more
open or deep water fish such as barracuda and tuna were exploited (Rose 1995; Shackleton
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1990; Stiner and Munro 2011). The subsistence picture from Franchthi Cave is that of
expanding diets, incorporating more low-ranked small game resources with time.
The Cave of Cyclope also has abundant Early and Late Mesolithic remains (Mylona
2003; Powell 2003; Trantalidou 2003). Ungulates, specifically goat and sheep, are common
throughout the sequence. A small number of pig remains are found in the later layers, and
land snails occur in the earlier layers (Trantalidou 2003). Abundant fish remains are found
throughout the Mesolithic but they increase in frequency in the Late Mesolithic (Mylona
2003; Powell 2003). As an island site marine resources are expected, but the Cave of
Cyclope confirms greater use of low-return small game, such as fish, in Mesolithic diets.
In Italy Middle Paleolithic hominins were successful hunters of large, prime-aged
adult ungulates in addition to opportunistic scavengers (Stiner 1994). The transport of
ungulate body parts is variable in the Italian Mousterian and seems to relate to specific food
acquisition tactics (e.g., hunting or scavenging) at different sites. Small game animals do
appear in the Middle Paleolithic assemblages, but they are generally restricted to easy to
collect prey such as tortoises and marine mollusks (Stiner 1994). In northern Italy, Middle
Paleolithic faunal assemblages are likewise dominated by prime-aged adult ungulate faunas,
though carnivores and small game such as lagomorphs and birds are also observed in low
numbers (Fiore, et al. 2004). The authors note that many of the bird and lagomorph remains
lack evidence of human modification, so it is unclear if they were introduced to the sites by
carnivores or other natural processes. A review by Alhaique et al. (2004) that spans the
Lower Paleolithic through Neolithic at sites around the country indicates that ungulates were
important throughout the Late Pleistocene and early Holocene, and that the frequencies of
small game increased in later periods.
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Ungulate exploitation continued in the Upper Paleolithic and Epipaleolithic at Italian
sites, as did small game collection, though the complexion of the small game component
changed (Stiner 2001; Stiner, et al. 1999; Stiner, et al. 2000). By the early Aurignacian, avian
faunas appear in the assemblages in considerable numbers, and by the Epipaleolithic
lagomorphs were heavily exploited. The inclusion of these small, fast-moving game species
was at the expense of slow-moving types, presumably because the latter were increasingly
rare in the environment. Declines in tortoise and shellfish in human diets may have been the
result of overharvesting; limpet diminution is apparent by 23,000 BP at Italian sites (Stiner
2001; Stiner, et al. 1999; Stiner, et al. 2000). However, shellfish gathering as a major food
source persisted at some Italian sites (Colonese, et al. 2011). In an analysis of late Upper
Paleolithic sites across Italy, Phoca-Cosmetatou (2003b) does not find an across-the-board
widening of diet breadth, but rather increasingly variable foraging patterns, one of which is
specialized ibex hunting. In northeast Italy, Phoca-Cosmetatou (2009) stresses the high
degree of variability in late Upper Paleolithic subsistence strategies and notes no specific
temporal trends in prey exploitation patterns if site use is not taken into account.
The French Paleolithic has been intensively studied and many high-quality data sets
are available. I limit my discussion here to those sites in the southern part of the country, in
the classic Mediterranean environmental regime (Figure 8.2). During the Middle Paleolithic
in southern France, hominins were highly mobile hunters that exclusively targeted large
ungulate prey (Boyle 2000; Valensi 2000). Boyle (2000) notes that regional variation in the
prey species exploited relates to geographic factors (i.e. montane regions typically associated
with ibex hunting). Species dominated by a single prey type are common, but may not reflect
specialized hunting reported elsewhere for the Upper Paleolithic. Boyle (2000) does not
331
exclude this possibility, but notes that seasonality data are currently lacking from many
Middle Paleolithic sites, so it is unclear if such monospecific faunas simply reflect season of
use. The author finds little evidence for scavenging in faunal assemblages from the region
based on the proportions of body parts found at Mousterian sites (Boyle 2000). At Lazaret
Cave, Valensi (2000) notes that Middle Paleolithic hominins hunted prime-aged adult red
deer and ibex, transporting the carcasses in whole form to the site for butchery.
Ungulate exploitation persisted in the Upper Paleolithic of Mediterranean France, and
species targeted largely reflect their environmental availability (Costamagno 2003; Surmely,
et al. 2003). The use of small game seems to correspond with the onset of the Upper
Paleolithic (Costamagno and Laroulandie 2004). Small animals occur in trace amounts at a
few Middle Paleolithic sites, though they may have been accumulated by carnivores. Small
game is likewise rare in the Châtelperronian but increases in frequency in later periods. By
the Magdalenian, the exploitation of birds, lagomorphs and fish is common (Costamagno and
Laroulandie 2004). Forager diets widen even further in the Mesolithic to include shellfish, in
addition to other small game resources.
Several faunal trends are apparent in Late Pleistocene sites of Mediterranean Spain.
Tortosa et al. (2002) provide an extensive review of Paleolithic faunal data from southern
Spain. They find that Middle Paleolithic hominins exploited a diverse range of ungulate
faunas, typically prime-aged adults or larger subadults (Tortosa, et al. 2002). Faunal richness
at Middle Paleolithic sites is often deceptively high because many human occupations,
particularly at rock shelters and caves, are interspersed with carnivore accumulations. Small
game is rare in the Middle Paleolithic, but recent evidence from Bolomor Cave indicates that
hominins at the site exploited tortoise, rabbit and duck as early as MIS 6 (Blasco 2008;
332
Blasco and Fernández Peris 2009; Sanchis Sera and Fernández Peris 2008). Species diversity
drops between the Middle and Upper Paleolithic as the result of contracting territories of
some herbivore species and the extinction of many large carnivores (Tortosa, et al. 2002).
Local fluctuations of ungulate taxa is probably in response to shifting environmental
conditions.
In the Upper Paleolithic, a range of small game animals appear in forager diets,
especially lagomorphs, but also including birds, fish and shellfish (Hockett and Haws 2002;
Tortosa, et al. 2002). During the Gravettian and Solutrian periods specialized deer, ibex and
horse hunting appears at certain sites. Adult and subadult ibex and red deer were targeted at
special-function sites in the Magdalenian and Epipaleolithic. In the later Upper Paleolithic
and Epipaleolithic, marine resources including mammals and shellfish appear in greater
numbers (Tortosa, et al. 2002).
It has been argued that many of the dietary differences between Middle and Upper
Paleolithic hominins in Spain stem from different land use strategies and increased
territoriality (Tortosa, et al. 2002; Villaverde, et al. 1996). During the Middle Paleolithic,
Neandertal groups had access to a greater range of large game resources, but by the Upper
Paleolithic, human foragers shifted between deer and ibex (i.e., coastal and mountainous
areas) on a seasonal basis. The authors conclude that rabbit exploitation is tied to this change
in land use and stress that it is not the result of human demographic pressure or resource
intensification (Tortosa, et al. 2002; Villaverde, et al. 1996). Whether or not this
interpretation is correct, the overarching trend fits with what is found in other parts of the
Mediterranean.
333
Also on the Iberian Peninsula is the Rock of Gibraltar, which has preserved a
concentration of large Paleolithic cave sites. Faunal data are available from the late
Mousterian layers from Vanguard Cave and the late Middle Paleolithic and post-LGM Upper
Paleolithic layers from Gorham’s Cave (Stringer, et al. 2008). The Vanguard Cave Middle
Paleolithic is dominated by marine mollusks and ungulates, specifically ibex and red deer,
though other ungulates are also found in addition to small amounts of carnivore, seal,
dolphin, bird, tortoise and fish (Stringer, et al. 2008). At Gorham’s Cave, lagomorphs are the
most common prey in both the Middle and Upper Paleolithic layers, followed by birds in the
Middle Paleolithic and cervids in the Upper Paleolithic, though again, other herbivores and
carnivores are also present in small numbers. Stringer et al. (2008) notes little change in the
faunal assemblages between the Middle and Upper Paleolithic layers, either in represented
species, prey diversity, or butchery patterns.
The Late Pleistocene faunal record of Portugal is less well-known than adjacent areas,
but recent work has greatly expanded our understanding of the western-most expanse of the
Mediterranean. One of the most striking features of the Portuguese record is the importance
of lagomorphs in the diet, at least during the Upper Paleolithic (Hockett and Haws 2002).
Middle Paleolithic faunal remains are fairly uncommon, but seem to focus on the
procurement of large game species (see Hockett and Haws 2009). Robust data sets are
available for some Upper Paleolithic sites, however.
Calderão Cave in north-central Portugal preserves Mousterian through Neolithic
remains, though the majority of the faunas associated with the Middle Paleolithic and Early
Upper Paleolithic layers were probably accumulated by hyenas (Davis 2002). In the later
Upper Paleolithic, a range of ungulates and carnivores occur in low frequencies, but the
334
proportion of rabbits is much greater (>95% in the Solutrean and Magdalenian layers) (Davis
2002).
At Vale Boi in Algarve, there is evidence of the intensive use of faunal resources
across the Upper Paleolithic. Limpets are abundant during the Gravettian but subsequently
decline during the Solutrian as the coastal geography changed (Manne and Bicho 2009).
Rabbits dominate the archaeofaunas in all Upper Paleolithic phases at the site, followed by
red deer and horse, which fluctuate in frequency probably as a result of climate shifts.
Significantly, there is extensive evidence for bone grease rendering, an extremely laborintensive activity, as early as the Gravettian (Manne and Bicho 2009; Manne, et al. 2005;
Stiner 2003b). Though no trends are evident in small game exploitation patterns or grease
rendering, it seems clear that by the Upper Paleolithic, human populations in southern
Portugal experienced some form of continuous resource stress.
Rabbits were likewise the main food source during the late Upper Paleolithic at
Picareiro Cave (Bicho, et al. 2006; Hockett and Bicho 2000). A range of ungulate species are
found at the site in low numbers, red deer and wild boar in particular, as well as small
carnivores, fish and shellfish. During the late Magdalenian, Bicho et al. (2006) argue that the
site was used as a hunting and carcass processing camp for small and large game, though
rabbit is by far the dominant species. The authors also suggest that there is evidence of
intensive marrow processing and bone grease rendering of large game animals at the site
(Bicho, et al. 2006). In Portugal, it seems that Upper Paleolithic subsistence strategies
differed from other areas of the Mediterranean, with the near-exclusive exploitation of lowranked prey and intensified processing of large game animals. On the other hand, quantitative
335
faunal data from the Middle Paleolithic and early Upper Paleolithic, which may indicate a
more widespread use of large game, are largely lacking at this point.
The main features of Late Pleistocene Mediterranean subsistence strategies are their
variability through space and time in response to local prey availability. Some broader trends
are apparent, however. Ungulate prey was preferred in all periods; evidence for the heavy
exploitation of small game typically corresponds with declines in large game hunting. Small,
sessile animals such as tortoises and marine mollusks are also exploited when they are
available in the environment. The timing of the inclusion of low-ranked prey such as small,
fast-moving hares, birds and fish varies regionally. In Iberia, for example, lagomorphs appear
in diets quiet early, possibly in the Middle Paleolithic but definitely with the Upper
Paleolithic. There are also some situations, such as when rabbit warrens are available, where
collecting young lagomorphs is argued to be less labor-intensive and does not require
specific technologies (Jones 2006). But the trend also involves hares in virtually every
Mediterranean region, and this species does not live underground. At the end of the
Pleistocene, diets included marine, terrestrial and avian faunas to a greater extent. In the most
general terms, then, there is a picture of intensification throughout the Mediterranean Basin
from the Late Pleistocene through the early Holocene, though understanding the dynamic
relations between local environmental factors and prey availability is critically important and
not always obvious.
Intensification and Specialization
There is the question of whether specialization and/or intensification occurred at
Klissoura Cave 1. These issues are often framed in terms of differences between the Middle
336
and Upper Paleolithic. Clearly, as discussed above, diet breadth widened across the
occupation sequence of the site and more low-ranked resources were included in the diet with
time, though mostly later in the Upper Paleolithic. In this sense, resource depression would
seem to have led to the intensified use of different species, particularly those taxa with high
fertility rates. On the other hand, there is no evidence of increased carcass processing in later
periods that would also be indicative of intensification. Perhaps resource depression
evidenced by changing prey types at the site was a local phenomenon, but comparisons to
other sites in southern Greece, including Franchthi Cave, would suggest otherwise. It
nonetheless is noteworthy that the evidence for foraging intensification does not include
increased marrow processing or bone grease rendering.
The dominance of fallow deer, particularly in the Middle Paleolithic layers of
Klissoura Cave 1 is also worth discussing. Some archaeologists have indicated that
specialized hunting was a hallmark of the Upper Paleolithic, at least in western Europe
(Costamagno 2003, 2004; Mellars 1973, 1989; 1991, but see Grayson and Delpech 2002,
2006). Indeed, specialized hunting as defined by monospecific assemblages is observed at
many Upper Paleolithic sites, including those with reindeer (David and Enloe 1993; Mellars
1989, 1991, 2004) and craggy-terrain-loving ibex and chamois (Gamble 1997; PhocaCosmetatou 2003a, 2003b, 2004a; Straus 1977, 1987). However, faunal assemblages
dominated by a single species are also found in the Middle Paleolithic (e.g., Adler, et al.
2006; Costamagno, et al. 2006; Gaudzinski 1995, 2006). The fundamental issue concerning
the question of specialization is what the existence of monospecific assemblages actually
means.
337
Monospecific or specialized assemblages contain a high proportion of a single taxon,
often as much as 90-95% of a single prey type (Mellars 1973, 1989, 2004), sometimes with
MNI values that approach nearly 100 animals (Gaudzinski 2006), though this is not a
requirement. From a taphonomic perspective, even if humans were responsible for
accumulating these faunas, several different hunting strategies can cause monospecific
assemblages, often as a palimpsest in the archaeological record. Gaudzinski (2006) notes a
few examples of different hunting strategies from the Middle Paleolithic: bovid-dominated
assemblages from the long-term exploitation of gregarious animals at Mauran, La Borde and
Wallertheim (Farizy, et al. 1994; Gaudzinski 1995; Jaubert, et al. 1990), the selection of
solitary young rhinoceros over a long period of time at Taubach (Bratlund 1999, though
rhinoceros only makes up 27% of this assemblage by NISP and 49% by MNI), and mass
hunting scenarios, such as reindeer at Salzgitter Lebenstedt (Gaudzinski and Roebroeks
2000) or Les Pradelles (Costamagno, et al. 2006). Clearly, there are many different situations
that result in monospecific assemblages.
The behavioral implications of specialization must be considered, and historically this
has been the source of many interpretive biases. Adler et al. (2006:90) rightly point out that
similarly diverse faunal assemblages are described as reflecting “opportunistic” hunting or
“increased diet breadth,” depending on whether they were accumulated by Neandertals or
modern humans, respectively, and Neandertal sites dominated by a single species have “low
diet breadth,” whereas a modern human monospecific assemblage indicates “specialization.”
For some authors, high proportions of a single species at a site may indicate intentional
selection of that species from all available prey (Mellars 1996), while others consider the
communal aspect of mass kills and processing (Binford 1968; David and Enloe 1993). Stiner
338
(1992:447) points out that specialization may simply reflect short-term adjustments to
seasonal or local food availability.
At Klissoura Cave 1, fallow deer dominate the faunal assemblages, comprising 6091% of the specimens identifiable to taxon in the Middle Paleolithic, and 12-78% in the
Upper Paleolithic and later layers. To be sure, there is considerable variation between the
assemblages within the Middle and Upper Paleolithic, but the larger point is that there is a
significant decline in fallow deer proportions over time. Fallow deer MNI values range
between one and fourteen individuals, depending on the layer. The proportions of fallow deer
are not over 90% except in the very earliest Middle Paleolithic layer. Based on the age
structure of individuals in the assemblages, this assemblage probably does not represent a
mass kill. The prevalence of fallow deer in most of the assemblages likely reflects a prime
fallow deer habitat in the Klissoura Gorge, particularly at the times of year when the site was
occupied the most frequently. Interestingly, fallow deer are not found to the southeast at
Franchthi Cave, though red deer are common, so perhaps the Klissoura Gorge remained
slightly drier and grassy during most periods.
CONCLUSION
The application of models from evolutionary ecology to understand the foraging
behaviors of hominins at Klissoura Cave 1 yields important conclusions about prey choice,
transport decisions, and butchering intensity and how these behaviors changed through the
sequence. Deviations from the modeled expectations are likewise interesting, such as the lack
of evidence for preferential transport of high-utility elements during periods when other lines
339
of evidence (e.g., hunting low ranked prey or increased marrow processing) indicate
intensification was occurring at the site.
Overall, trends in faunal exploitation at Klissoura Cave 1 fit fairly well with those
found at other Late Pleistocene sites in the Mediterranean, but there are also differences
brought on by factors of the local environment and unique cultural adaptations in Greece.
The presence of a long, well-preserved sequence with Middle Paleolithic, Uluzzian, abundant
Aurignacian and overlying Upper Paleolithic and Mesolithic deposits is unprecedented for
southern Greece. One of the more striking features of the long faunal sequence at Klissoura
Cave 1 is the relatively static nature of the Middle Paleolithic record, as compared to the
highly variable Upper Paleolithic hunting, transport, and carcass butchery strategies. Trends
in the latter largely reflect changes in site use and occupation intensity. Overarching trends
through the sequence concern mainly the increased importance of small game, particularly
fast-moving animals, with time, which begins as early as the Aurignacian. Ungulates
dominate the assemblages until the Epigravettian and Mesolithic layers, when small game
overtakes large taxa by NISP counts. This tendency generally parallels the situation in many
parts of the Mediterranean, including other sites in Greece, where Mesolithic foragers turned
their attention to low-ranked resources such as birds, hares, and especially fish. Human
groups at Klissoura Cave1 did not uniformly contend with resource shortages by intensifying
carcass processing, though this did occur elsewhere in the Mediterranean Basin (e.g.,
Portugal and the Levant), but the occupants of Klissoura Cave 1 did intensify marrow
processing efforts in the Aurignacian (layer IV). It is possible that intensification strategies
such as bone grease rendering never entered the repertoire of foragers in southern Greece, but
340
it should also be noted that grease rendering appeared elsewhere in Europe after the
Aurignacian and therefore after the heaviest use of Klissoura Cave 1.
Klissoura Cave 1 preserves a long sequence, and its location provides the opportunity
to address two of the major geographical gradients in Mediterranean Europe. First, are eastwest trends in species availability and the expansion and establishment of modern human
populations, and subsequent cultural diversification during the early Upper Paleolithic.
Second are the ecological conditions or refugium status of one of three major European
peninsulas (Iberia, Italy, and southern Greece) during the climatically variable Pleistocene.
Faunal evidence indicates that Klissoura Cave 1 was used mostly in the winter, particularly
during the Middle Paleolithic, possibly following ungulates into lowland pastures with milder
temperatures. Assuming modern humans were the manufacturers of Aurignacian stone tool
industries, there is no evidence to suggest that Neandertal populations persisted later than
40,000 BP in southern Greece, as opposed to the situation in Iberia. Rather, there is a spike in
human activity at Klissoura Cave 1 not long after modern human populations are expected to
have appeared in southern Greece. Site use dropped significantly after the early Upper
Paleolithic. The faunal trends at Klissoura Cave 1 nonetheless suggest increases in human
populations locally and regionally, and the taxing of high-value faunal resources. This caused
foragers in southern Greece to turn to lower-ranked terrestrial and marine resources by the
end of the Upper Paleolithic and Mesolithic, before the eventual adoption of agricultural
lifeways with the onset of the Neolithic.
341
APPENDICES
Appendix A: Hare survivorship and bone density values (values from Pavao and Stahl 1999).
342
Appendix B: Fallow deer survivorship and bone density values (values from Lyman 1994).
343
Appendix B - continued
344
Appendix B - continued
345
Appendix B - continued
346
Appendix C: Tooth and bone-based MNE values by layer.
347
Appendix D: Shaft and end-based MNE values by layer.
348
Appendix E: Middle Paleolithic hare elements. Horizontal line indicates lowest structural density for
cervids.
349
Appendix F: NISP and MNE values by layer for all taxon.
350
Appendix F - continued
351
Appendix G: Biomass values for common taxa.
352
Appendix G - continued
353
Appendix G - continued
354
Appendix H: Utility indices for Rangifer, combined by anatomical region (values from Binford
1978).
355
Appendix I: Percent MAU for fallow deer by layer.
356
Appendix I - continued
357
Appendix I - continued
358
Appendix J: Mean FUI for fallow deer by layer.
359
Appendix J - continued
360
Appendix K: Fallow deer anatomical representation.
361
Appendix K - continued
362
Appendix L: Hare anatomical representation.
363
Appendix M: Partridge anatomical representation.
364
Appendix N: Anatomical representation for red deer and ibex in layer IV.
365
Appendix O: Tooth wear stages for ungulates by layer.
366
Appendix O - continued
367
Appendix P: Ungulate fusion stages (values from Reitz and Wing 2008 and Silver 1969).
368
Appendix P - continued
369
Appendix P - continued
370
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