Fishing, Curing and Smoking Fish at Cueva De Los Vampiros: a Contextual and Archeofaunal Evaluation of a Purported Pre-Columbian

UNIVERSITY OF CALGARY

Fishing, curing and smoking fish at Cueva de los Vampiros: a contextual and archeofaunal

evaluation of a purported Pre-Columbian fishing Camp near Parita Bay (Panama, Central

Pacific).

Diana Rocío Carvajal Contreras

A THESIS

SUBMITTED TO THE FACULTY OF GRADUATE STUDIES

IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE

DEGREE OF DOCTOR OF PHILOSOPHY

DEPARTMENT OF ARCHAEOLOGY

CALGARY, ALBERTA

2010

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Abstract

This doctoral dissertation is a study of fish remains from a site on the Pacific coast of Panama. The objective is to determine through a study of the fish bones whether fish were subjected to preservative technologies at the site and whether the site may have served as specialized location to prepare fish to transport to settlements situated further inland.

Fish and fish products are reported in ethnohistoric documents of Panama as important trade commodities among the chiefdoms of the region. Two rockshelters at Cueva de los Vampiros show evidence for having been used intensively for fishing and preparing fish between ca. 2200 and 1900 BP. Evidence is presented of human impacts on fish skeletons resulting from the in situ preparation of fish (i.e., gutting, cutting and smoking) providing the opportunity to compare the archaeofaunal data with two other sites, Sitio Sierra and AG-125, and with ethnoarchaeological work conducted by Irit Zohar.

Several lines of evidence: zooarchaeological (vertebrate and invertebrate remains), taphonomical (butchering patterns, postdepositional events), anthropic features (postholes, pits and hearths) and artefactual (pottery, lithic and shell tools) data, which in conjunction; support the hypothesis that Vampiros shelters were a place where Pre-Columbian inhabitants brought large numers of captured fish and prepared them for transport elsewhere. The high frequency of puffer fish here, in contrast with other sites in Central Pacific Panama, suggests that Pre-Columbian inhabitants at Vampiros were perhaps removing the skin, guts, tail and head of puffer fish animals and popping-out two dorsal fillets and exporting the fillets as a delicacy to Sitio Sierra during the dry season.

Non fish vertebrate remains indicate that Vampiros inhabitants were not only exploiting and processing fish and gathering molluscs but they were engaged in other local activities in a highly modified coastal landscape.

The evidence is insufficient to evaluate conclusively the role of the Vampiros shelters in the local and regional economy during the Middle Ceramic Period to provision inland sites like Sitio Sierra with inshore marine fish. However, it does demonstrate that the people of Panama, as early as early as 2200 BP, had developed techniques for preserving fish— techniques that later would be critical for thedevelop of regional trade in fish among the chiefdoms of Panama.

Key Words

Zooarchaeology, Fish, Mollusks, Taphonomy, Intermediate Area, Panama, Coastal Resources, Neotropics

iii

Acknowledgements

Thanks to everyone who helped me with the completition of this project. First and foremost

I would like to thank the member of my dissertation committee for their useful comments Dr

Scott Raymond, Dr Brian Kooyman, Dr Geoffrey McCafferty, Dr Denise Brown (University of Calgary) and Dr Anthony Ranere (Temple University). The fieldwork and laboratory work of this investigation was possible thanks to a Smithsonian ten-week graduate fellowship and

Smithsonian Pre-doctoral fellowship and the University of Calgary´s James Carter fund.

Financial support for my doctoral studies was provided through a University of Calgary

Graduate Research and Dean´s Entry Scholarships and Nicholls International Graduate

Archaeology Scholarship.

Permission to conduct fieldwork and to transport fish samples to Canada was provided by

Department of Patrimonio Histórico at Instituto Nacional de Cultura (Panama) during my research.

I owe a tremendous debt of gratitude to two persons: my supervisor Dr Scott Raymond for his counsel throughout my doctoral studies, his comments on numerous course papers, articles and drafts of the dissertation and Dr Richard Cooke who acted as my advisor during my research at the Smithsonian Tropical Research in Panama. He opened his laboratory, gave me Sitio Sierra and Ag-125 fish samples, trained me in faunal analysis and provided advice and comments throught this research. The radiocarbon dates to this project were iv

funded by an annual research allowment made by the Smithsonian Institution to Dr Richard

Cooke.

I would like to thank everyone at Smithsonial Tropical Research Institute, in particular

Máximo Jiménez, Alexandra Lara Kraudy, Aureliano Valencia, Lisbeth Valencia, Jaqueline

Sanchez, Conrado Tapia, Felix Rodriguez and Adriana Bilgray for their assistance during my research and laboratory work.

During the course of this dissertation, several people gave useful comments, theses and articles, particulary Dr Georges Pearson, Dr Irit Zohar, Dr Patricia Hansell, Dr Phillipe

Bearez, Dr Anne Katzenberg, Dr Gerry Oetelaar, Dr German Peña, Pedro Jose Botero, Isabel

Rivera M.A and Dr Christopher Gotz.

To all those who assisted in the field in Panama, I express my sincere appreciation. I am particularly indebted to Ubencio Vargas, Lisbeth Valencia and Lourdes Castillero who worked with me as assistants. Others in Panama who deserve thanks are Luis Sánchez, for his contributions of drawings of pottery, and Dr Ilean Isaza, who provided information about

Central Pacific Panama archaeology.

I am extremely grateful to many people including Dr Jane Kelley, Dr. Len Hills, Dr. Latonia

Hartery, Dr Julia Mayo, Larry Steinbrenner M.A, Laura Roskowski M.A, Ilicena Tapia,

Nicole Ethier, Lilly Wong, Kitty Raymond, Angélica Lopez-Forment, Manuel Arturo

Izquierdo, Fernando Astudillo, Alejandra Alonso, Sergio Gaytan, Anastasia Antonova,

Shawn Morton, Meaghan Peuramaki- Brown, Guncha Byashiyeva, Marisha Shegda, Deepika

Fernandez, Janet Blakey, Julie-Anne White, Juan Guillermo Martin, Paula Figueroa, and v

Claudia Valderrama that made my stay in Panama and Canada confortable and enjoyable, and provided me with editorial comments, help, food, friendship and moral support.

Finalmente y no por eso menos importante quiero agradecer con besos y abrazos a mi familia: mis padres Gloria y Nelson a quienes dedico esta tesis y mis hermanos Diego y Eduardo.

Todos ellos me empujaron, animaron y apoyaron de todas las formas posibles a lo largo de esta tesis y mi carrera profesional.

Table of Contents

Approval Page ...... i Abstract ...... iii Acknowledgements ...... iv Table of Contents ...... vii List of Tables ...... x List of Figures and Illustrations ...... xvii List of Symbols, Abbreviations and Nomenclature ...... xxviiii Appendices...... xxix

CHAPTER 1:INTRODUCTION ...... …………1 Coastal Resources ...... 2 Fish and Trade ...... 6 Objectives ...... 10

CHAPTER 2: CUEVA DE LOS VAMPIROS SITE (AG-145) ...... 14 Introduction ...... 14 Modern Environmental Setting ...... 16 Paleo-environmental Setting ...... 20 Vampiros Rockshelters Background Considerations ...... 26 Stratigraphy and Deposits ...... 33 Macrounit 1 and 2 ...... 38 Field Methods ...... 47

CHAPTER 3: CUEVA DE LOS VAMPIROS AND ITS CULTURAL-GEOGRAPHICAL SETTING ...... 49 Introduction ...... 49 Central Pacific Panama ...... 52 Chronology ...... 54 Fishing in Central Panama………………………………………………………………..57 Ethnohistorical Evidence of Exchange and Curing Fish in Central Panama……………...65

CHAPTER 4: POTTERY, SHELL AND LITHIC ARTIFACTS AT CUEVA DE LOS VAMPIROS SITE (AG-145) ...... 70 Introduction ...... 70 Stone Tools ...... 70 Chipped Stone Tools ...... 72 vii

Ground and Polished Stone Tools ...... 75 Possible Function of the Stone tools……………………………………………………..77 Shell Ornaments ...... 78 Pottery ...... 79 Pottery Sherds and Site Formation ...... 84 Summary ...... 88

CHAPTER 5: MOLLUSC REMAINS AT CUEVA DE LOS VAMPIROS SITE (AG-145)89 Introduction ...... 89 Habitats Used for Shellfish Gathering ...... 91 Species Selection ...... 98 Dominance and Contribution to Diet ...... 100 Shellfish size and Gathering Methods ...... 110 State of Preservation, Human and Natural Modifications……………………………….122 Interpretation ...... 136

CHAPTER 6: OTHER VERTEBRATE REMAINS AT CUEVA DE LOS VAMPIROS SITE (AG-145) ...... 141 Introduction ...... 141 Analytical Procedures ...... 141 Results ...... 142 Macrounit 1 ...... 143 Macrounit 2a ...... 156 Macrounit 2b ...... 163 Discussion ...... 168

CHAPTER 7: PRECOLUMBIAN FISHING AT CUEVA DE LOS VAMPIROS SITE (AG-145) ...... 170 Introduction ...... 170 Recovery Methods and Sample bias ...... 170 Identification and Comparative Collection ...... 171 Quantification Analysis ...... 172 Taphonomic Analysis ...... 177 Discussion of Element Distribution, Fractures and Cut Marks from Various Fish Butchering Processes...…………………………………………………………………180 Fish Families and Environment ...... 188 Diversity and Equitability...... 201 Dominance and Contribution to Diet ...... 203

viii

Natural Transformation Processes and Bones ...... 221 Cultural Processes and Bones ...... 225 Relative Representation of Skeletal Elements ...... 228 Articulated bones ...... 250 Damage………………………………………………………………………...………257 Cutmarks ...... 259 Pattern of Fractures ...... 266 Summary ...... 277

CHAPTER 8: PRECOLUMBIAN FISHING AT OTHER SITES ON THE SANTA MARIA´S WATERSHED ...... 287 Introduction ...... 287 Background of Comparative Samples ...... 288 Methodology ...... 289 Sitio Sierra´s Sample (AG-3) ...... 290 Results ...... 294 AG-125‘s Sample ...... 307 Results ...... 310 Butchering Units or Articulated Bones ...... 324 Cutmarks ...... 326 Damage………………………………………………………………………………...329 Fractures...... 331 Summary ...... 345

CHAPTER 9: DISCUSSION AND CONCLUSIONS ...... 356 Introduction ...... 356 Paleoenvironemtal Interpretation ...... 357 Exploitation of Terrestrial and Aquatic Resources....………….…..………………….…367 Taphonomy and butchering………………………………………………………….....370 Exchange of Fish and Specialization in Panama as seen from Vampiros Rockshelters……………………………………………………………………………380 Comparisons…………………………………………………………………….……...386 Conclusions and Future Research………………………...…………………………….390

REFERENCES ...... …………………………………………………. 393

List of Tables

Table 2.1: Soil characteristics of the stratigraphic subunits at Macrounit 1...... 38

Table 2.2: Soil characteristics of the stratigraphic subunits at Macrounit 2a...... 40

Table 2.3: Soil characteristics of the stratigraphic subunits at Macrounit 2b...... 42

Table 3.1: Chronological sequence of ―Gran Coclé‖...... 54

Table 3.2: Archaeological sites in Central Panama: distance from the sea, dates, animal resources and environment...... 61

Table 3.3: Archaeological sites in Central Panama: methods of fishing and fishing artefacts.63

Table 4.1: Types of Stone Tools at Vampiros Rockshelters……………………………...... 70

Table 4.2: Types of materials at Vampiros Rockshelters ...... …………………………….71

Table 4.3: Type of Flakes at Vampiros Rockshelters ...... 72

Table 4.4: Flakes: Type of Impact ...... 72

Table 4.5: Type of Ground/Polished Materials at Vampiros Rockshelters ...... 76

Table 4.6: Radiocarbon Dates at Vampiros...... 81

Table 5.1: Mollusc diversity and Equitability at Vampiros-1...... 99

Table 5.2: Mollusc diversity and Equitability at Vampiros-2...... 99

Table 5.3: Bivalve assemblage at Vampiros-1...... 104

Table 5.4: Gastropod assemblage at Vampiros-1 ...... 106

Table 5.5: Bivalve assemblage at Vampiros-2 ...... 108

Table 5.6: Gastropod assemblage at Vampiros-2………………………………………….110

Table 6.1: Non- fish vertebrate class frequencies from Vampiros-1 and Vampiros- 2 expressed in number of identifiable specimens (NISP and NISP%)……………………….142

Table 6.2: Fish versus Non- fish vertebrate from Vampiros-1 and Vampiros- 2 expressed in number of identifiable specimens (NISP and NISP %)……………………………………143

Table 6.3: Reptiles at macrounit 1(NISP, MNI and estimated biomass)…………………...143

Table 6.4: Reptiles at macrounit 1: modifications (NISP)…………………………………145

Table 6.5: Mammals macrounit 1 (NISP, MNI and estimated biomass)…………………....148

Table 6.6: Mammals at macrounit 1: modifications (NISP)………………………………...150

Table 6.7: Amphibia at macrounit 1 (NISP, MNI and estimated biomass)...... 153

Table 6.8: Amphibian at macrounit 1: modifications (NISP)……………………………...153

Table 6.9: Aves at macrounit 1 (NISP, MNI and estimated biomass)...... 155

Table 6.10: Birds at macrounit 1: modifications (NISP)…………………………………...155

Table 6.11: Reptiles at macrounit 2a (NISP, MNI and estimated biomass) ...... …….156

Table 6.12: Reptiles at macrounit 2a: modifications (NISP)………………………………159

Table 6.13: Mammals at macrounit 2a (NISP, MNI and estimated biomass) ...... 161

Table 6.14: Mammals at macrounit 2a: modifications (NISP)……………………….……..161

Table 6.15: Aves at macrounit 2a (NISP, MNI and estimated biomass) ...... 162

Table 6.16. Birds at macrounit 2a: modifications (NISP)…………………………………..162

Table 6.17: Reptiles at macrounit 2b (NISP, MNI and estimated biomass)...... 163

Table 6.18: Reptiles at macrounit 2b: modifications (NISP)………………………………164

Table 6.19: Mammals at macrounit 2b (NISP, MNI and estimated biomass) ...... …..166

Table 6.20: Mammals at macrounit 2b: modifications (NISP)……………………………...167

Table 6.21: Aves at macrounit 2b (NISP, MNI and estimated biomass) ...... 167

Table 7.1: Butchering methods describe by Zohar (2003), Zohar and Cooke (1997) and Zohar et al (2001)...... 186

Table 7.2: Distintion between predepositional and postdepositional...... 187

Table 7.3: Criteria for identying cutmarks and fractures...... 188

Table 7.4: (NISP) and percentage of fish remains identified and unidentified by loci……...189

Table 7.5 Complete analyzed units: (NISP) and percentage of identified fish remains by mesh, shelter and macrounit including the rank (in red) most abundant families…………………190

Table 7.6: Supplementary analyzed units: (NISP) and percentage of identified fish remains by mesh, shelter and macrounit………………………………………………………………191

Table 7.7: The ecological composition of the most abundant species and the probable method of fishing………………………………………………………………………….201

Table 7.8: Diversity an equitability at Vampiros 1 and 2 by MNI………………………….203

Table 7.9: Diversity an equitability at Vampiros 1 and 2 by biomass ...... 203

Table 7.10:Vampiros 1 and 2: Fish species list at Macrounit 1…………………………...... 205

Table 7.11: Vampiros 1 and 2 : Fish species list at Macrounit 2a…………………………210

Table 7.12: Vampiros 1 and 2: Fish species list at Macrounit 2b…………………………..216

Table 7.13: Summary articulated bones(NISP) at Vampiros 1 and 2 by mesh and macrounit...... 222 Table 7.14: Summary natural processes on fish bones (NISP) at Vampiros 1 and 2) by mesh and macrounit…………………………………………………………………………….224

Table 7.15: (NISP) and percentage of burnt fish remains identified and unidentified by loci and mesh (fine mesh including 1/16‖ and 0.25)...... 227

Table 7.16: Differential heat exposure, (NISP) and percentage of burnt fish remains from complete analyzed units by macrounit……………………………………………………..227

Table 7.17: Articulated bones (NISP) at Vampiros 1 and 2 by macrounit...... 250

Table 7.18: Macrounit 2, Family Ariidae: articulated bones by anatomic region…………..252

Table 7.19: Macrounit 2, Family Carangidae: articulated bones by anatomic region ...... …..252

Table 7.20: Macrounit 2, Families Albulidae, Belonidae, Clupeidae, Polynemidae, Pristigasteridae, Sciaenidae and Scombridae: articulated bones by anatomic region………. 253

Table 7.21: Macrounit 2, Family Tetraodontidae: articulated bones by anatomic region...... 253

Table 7.22: Vampiros 1 and 2: cur marks by macrounit and mesh size( I= Indeterminate, P= Predepositional, S=Postdepositional)……………………………………………………...258

Table 7.23: Summary distribution of cut marks atMacrounit 2,Ariidae family...... 259

Table 7.24: Distribution of predepostional cut marks at Macrounit 2, Ariidae family……..260

xii

Table 7.25: Distribution of predepostional cut marks at Macrounit 2,Belonidae family.…..261

Table 7.26: Distribution of predepostional cut marks at Macrounit 2,Carangidae family….261

Table 7.27: Distribution of predepostional cut marks at Macrounit 2, Haemulidae family.262

Table 7.28: Distribution of predepostional cut marks at Macrounit 2, Pristigasteridae family………………………………………………………………………………...... 262

Table 7.29: Summary distribution of cut marks at Macrounit 2, Sciaenidae family...... 263

Table 7.30: Distribution of predepostional cut marks at Macrounit 2,Sciaenidae family…...263

Table 7.31: Distribution of predepostional cut marks at Macrounit 2, Tetraodontidae family……………………………………………………………………………………...264

Table 7.32: Distribution of predepostional cut marks at Macrounit 2, Albulidae, Batrachoididae, Centropomidae, Lobotidae, Lutjanidae family…………………………….264

Table 7.33: General frequency (NISP) and percentage of fragmentation at Vampiros….…266

Table 7.34: Frequency (NISP) and percentage of fragmentation at Vampiros in completed analyzed samples by macrounit…………………………………………………………….266

Table 7.35: Frequency (NISP) and percentage of predepositional and postdepositional fractures at Vampiros in completed analyzed samples by macrounit……………………….267

Table 7.36: Frequency (%) of Ariidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Vampiros (macrounit 2)………………………………………………………………………………………….268

Table 7.37: Frequency (%) of Belonidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Vampiros (macrounit 2) ……………………………………………………………………………………………268

Table 7.38: Frequency (%) of Clupeidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Vampiros (macrounit 2)………………………………………………………………………………………….269

Table 7.39: Frequency (%) of Carangidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Vampiros (macrounit 2) ……………………………………………………………………………………………269

Table 7.40: Frequency (%) of Haemulidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Vampiros (macrounit 2) …………………………………………………………………………….270 xiii

Table 7.41: Frequency (%) of Polynemidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Vampiros (macrounit 2)……………………………………………………………………………...271

Table 7.42: Frequency (%) of Pristigasteridae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Vampiros (macrounit 2) ……………………………………………………………………………..271

Table 7.43: Frequency (%) of Sciaenidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Vampiros (macrounit 2)…………………………………………………………………………………………..272

Table 7.44: Frequency (%) of Tetraodontidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Vampiros (macrounit 2)……………………………………………………………………………...272

Table 7.45: Weighted mean index (WMI) of highly fragmented bones and portion (NISP) for Ariidae and Belonidae in Vampiros (macrounit 2) ………………………………………..273

Table 7.46: Weighted mean index (WMI) of highly fragmented bones and portion (NISP) for Carangidae, Clupeidae and Haemulidae in Vampiros (macrounit 2) ………………………274

Table 7.47: Weighted mean index (WMI) of highly fragmented bones and portion (NISP) for Polynemidae, Pristigasteridae and Sciaenidae in Vampiros (macrounit 2) …………………274

Table 7.48: Weighted mean index (WMI) of highly fragmented bones and portion (NISP) for Tetraodontidae in Vampiros (macrounit 2)………………………………………………..275

Table 7.49: Types of fractures observed on the cleithrum on the most abundant families at Vampiros………………………………………………………………………………….276

Table 8.1: Sitio Sierra: Fish species list…………………………………………………..…291

Table 8.2: Differential heat exposure (NISP) and percentage of burnt fish remains from Sitio Sierra………………………………………………………………………………………294

Table 8.3: Ag-125: Fish species list……………………………………………………..…308

Table 8.4: Differential heat exposure, (NISP) and percentage of burnt fish remains from Ag- 125………………………………………………………………………………………...311

Table 8.5: Sitio Sierra: articulated bones by anatomic region………………………………319

Table 8.6: Distribution of cut marks at Sitio Sierra and Ag-125……………………………328

Table 8.7: Frequency (NISP) and percentage of predepositional and postdepositional fractures and completeness at Sitio Sierra…………………………………………………………...330

xiv

Table 8.8: Frequency (NISP) and percentage of fragmentation at Sitio Sierra……………...330

Table 8.9: Frequency (%) of Ariidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Sitio Sierra……………331

Table 8.10: Frequency (%) of Carangidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Sitio Sierra……………332

Table 8.11: Frequency (%) of Clupeidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Sitio Sierra……………333

Table 8.12: Frequency (%) of Haemulidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Sitio Sierra……334

Table 8.13: Frequency (%) of Sciaenidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Sitio Sierra…………..335

Table 8.14: Weighted mean index (WMI) of highly fragmented bones and portion (NISP) for Sitio Sierra sample………………………………………………………………………..336

Table 8.15: Types of fractures observed on the cleithrum on the most abundant families at Sitio Sierra………………………………………………………………………………..337

Table 8.16: Frequency (NISP) and percentage of predepositional and postdepositional fractures and completeness at Ag-125……………………………………………………338

Table 8.17: Frequency (NISP) and percentage of fragmentation at Ag-125……………….338

Table 8.18: Frequency (%) of Ariidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Ag-125………………339

Table 8.19: Frequency (%) of Carangidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Ag-125……………….340

Table 8.20: Frequency (%) of Clupeidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Ag-125……………….341

Table 8.21: Frequency (%) of Haemulidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Ag-125……...…342

Table 8.22: Frequency (%) of Sciaenidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Ag-…………………..344

Table 8.23: Frequency (%) of Tetraodontidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Ag-125………..343

Table 8.24: Weighted mean index (WMI) of highly fragmented bones and portion (NISP) for Ag-125 sample……………………………………………………………………………344 xv

Table 9.1: Table listing some aquatic and terrestrial taxa divided by site, date, distance from the sea, habitat, probable method of fishing and some fishing artefacts at Middle ceramic sites in Central Pacific Panama...... 364

Table 9.2: Table comparing biomass in some aquatic and terrestrial animals at Vampiros by macrounit...... 365

Table 9.3: Summary of Vampiros, Sitio Sierra and Ag-125...... 382

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List of Figures and Illustrations

Figure 1.1: Map of the study area ...... 2

Figure 2.1: Map and aerial photography showing the location of the Vampiros-1 and Vampiros-2 shelters with respect to other pre-Columbian sites around Parita...... ………………...... 15

Figure 2.2: Map of the study area showing environmental setting surrounding Vampiros rockshelters (AG-145), Sitio Sierra (AG-308) and AG-125...... 19

Figure 2.3: Map of Panama´s coastlines during the Pleistocene ...... 21

Figure 2.4: Hypothetic Parita Bay´s ancient coastline during 7000 BP…………….……….23

Figure 2.5: Hypothetic Parita Bay´s ancient coastline during 2200 BP ...... 25

Figure 2.6: Hypothetic Parita Bay´s ancient coastline during 1100 BP ...... 26

Figure 2.7: Location of Vampiros-1. Top: Cross section of Vampiros-1 (Pearson 2002). Left: View of the east wall with the 2005 columns. Right: General view (Pearson 2002)...... 29

Figure 2.8: Map of Vampiros-1 showing excavations units 1982-2006 (TP) and column samples 2005 #1 (right) and # 2 (left) ...... 30

Figure 2.9: Location of Vampiros-2.Top: Cross section of Vampiros-2 (Pearson). Left: View of the north wall with the 2005 column. Right: General view (Pearson) ...... 31

Figure 2.10: Map of Vampiros-2 showing excavations units 2005-2006 (TP) and the 2005 column sample# 3 ...... 32

Figure 2.11: Profiles of part of the south and the west walls at Vampiros-1 showing the relationship between depositional units (D) 1 and 2 (‗‗fishing camps‘‘) and D 3–5 (Drawing: G.A. Pearson) (Carvajal Contreras et al 2008: 98)…………………………………………..33.

Figure 2.12: Stratigraphic profile of Macrounit 1 and 2, Vampiros-2 (north wall) and Vampiros-1 (east wall)...... 36

Figure 2.13: Radiocarbon dates for Vampiros-1 and Vampiros-2……………………..……..37

Figure 2.14: Detailed stratigraphic profile of column 1, Vampiros-1...... 39

Figure 2.15: Detailed stratigraphic profile of column #2, Vampiros-1……………………...41

xvii

Figure 2.16: Detailed stratigraphic profile of column #3, Vampiros-2 ...... ……43

Figure 2.17: Geochemical analyses of Vampiros-1 (top) and Vampiros-2 (bottom)………..44

Figure 3.1: Panama´s map including archaeological sites…………………………………..51

Figure 3.2: Native Americans transporting goods and crossing a river in Costa Rica ...... 66

Figure 4.1: Left, tertiary flake with use-wear. Right, Flake from a celt ...... 73

Figure 4.2: Left, La Mula point made of yellow jasper, Right, La Mula point made of unknown material...... 74

Figure 4.3: Left, Blade fragment. Center, blade with use- wear. Right, tertiary flake with potlids scars………………………………………………………………………………...74

Figure 4.4: Unifacial scraper ……………………………………………………………….75

Figure 4.5: Celt found at Vampiros-1……………………………………………………….76

Figure 4.6: Unmodified pebbles……………………………………………………………77

Figure 4.7: Shell ornament found at Vampiros-1……………………………………………79

Figure 4.8: Comparison painted sherds (Las Huacas, La Mula-Sarigua, Sitio Sierra, Vampiros…………………………………………………………………………………...82

Figure 4.9: Comparison plastic decorations Cerro Juan Diaz, La Mula- Sarigua, Cueva de los Vampiros…………………………………………………………………………………...83

Figure 4.10: Top: Contour drawing refitting at Vampiros-1. Bottom: Mesh drawing refitting zones at Vampiros-1………………………………………………………...... ……...86

Figure 4.11: Contour drawing refitting at Vampiros-2 Bottom: Mesh drawing refitting zones at Vampiros-2…………………………………………………………………………………87

Figure 5.1: Percentage of mollusks remains in macrounit 1 at Vampiros-1 and Vampiros-2 by weight………………………………………………………………………………………92

Figure 5.2: Percentage of mollusks remains in macrounit 2a at Vampiros-1 and Vampiros-2 by weight………………………………………………………………………………………93

Figure 5.3: Percentage of mollusks remains in macrounit 2b at Vampiros-1 and Vampiros-2 by weight……………………………………………………………………………………....94

Figure 5.4: Exploited habitats at Vampiros-1 (MNI)………………………………………..96

Figure 5.5: Exploited habitats at Vampiros-2 (MNI)………………………………………..97 xviii

Figure 5.6: Argopecten ventricosa´s boxplots. The line indicates Keen´s average length and then number enclosed on the box is the MNI……………………………………………..112

Figure 5.7: Polinices otis box plots. The line indicates Keen´s average height and then number enclosed on the box is the MNI…………………………………………………………..113

Figure 5.8: Pitar paytensis box plots. The line indicates Keen´s average length and then number enclosed on the box is the MNI………………………………………………….114

Figure 5.9: Natica unifasciata box plots. The line indicates Keen´s average height and then number enclosed on the box is the MNI…………………………………………………..115

Figure 5.10: Grandiarca grandis box plots. The line indicates Keen´s average length and then number enclosed on the box is the MNI……………………………………………….…116

Figure 5.11: Thais kiosquiformis box plots. The line indicates Keen´s average height and then number enclosed on the box is the MNI…………………………………………………..117

Figure 5.12: Protothaca asperrima box plots. The line indicates Keen´s average length and then number enclosed on the box is the MNI…………………………………………….118

Figure 5.13: Anadara tuberculosa box plots. The line indicates Keen´s average length and then number enclosed on the box is the MNI…………………………………………………..119

Figure 5.14: Tellina laceridens box plots. The line indicates Keen´s average length and then number enclosed on the box is the MNI…………………………………………………..120

Figure 5.15: Dosinia dunkeri box plots. The line indicates Keen´s average length and then number enclosed on the box is the MNI………………………………………………..…121

Figure 5.16: Mollusk remains with abrasion per subunit in each macrounit and shelter by weight…………………………………………………………………………………...…123

Figure 5.17: Mollusk remains with acid dissolution per subunit in each macrounit and shelter by weight…………………………………………………………………………………..124

Figure 5.18: Mollusk remains with encrustation per subunit in each macrounit and shelter by weight……………………………………………………………………………………..125

Figure 5.19: Bivalve remains completeness (fragment, semi-fragment, whole) per subunit in each macrounit and shelter by weight…………………………………………………...…127

Figure 5.20: Gastropod remains completeness (fragment, semi-fragment, whole) per subunit in each macrounit and shelter by weight ……………………………………………………..128

Figure 5.21: Mollusc remains heat treatment (black, grey, brown) per subunit in each macrounit and shelter by weight…………………………………………………………..129

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Figure 5.22: Paired molluscan remains per subunit in each macrounit and shelter by weight……………………………………………………………………………………..130

Figure 5.23: Mollusc remains with natural perforation per subunit in each macrounit and shelter by weight…………………………………………………………………………...131

Figure 5.24: Completeness ranges among taxa at Vampiros-1……………………………...132

Figure 5.25: Completeness ranges among taxa at Vampiros-2…………………………...... 133

Figure 5.26: Heat treatment among taxa at Vampiros-1……………………………………134

Figure 5.27: Heat treatment among taxa at Vampiros-2……………………………………135

Figure 6.1: Element distribution of freshwater turtles expressed in number of identified specimens (NISP)………………………………………………………………………….144

Figure 6.2: Element distribution of marine turtles expressed in number of identified specimens (NISP)……………………………………………………………………………………146

Figure 6.3: Element distribution of Iguana expressed in number of identified specimens (NISP)…………………………………………………………………………………….147

Figure 6.4: Element distribution of small mammals expressed in number of identified specimens (NISP)………………………………………………………………………….149

Figure 6.5: Mammal and reptile bones and teeth from the Vampiros shelters, Cerro Tigre, Cocle, Panama (Photos: Tara Hornung)…………………………………………………...151

Figure 6.6: Mammal, bird and reptile bones from the Vampiros shelters, Cerro Tigre, Cocle´, Panama (Photos: Tara Hornung)………………………………………………………….152

Figure 6.8: Element distribution of freshwater turtles expressed in number of identified specimens (NISP)…………………………………………………………………………157

Figure 6.9: Element distribution of Iguanas expressed in number of identified specimens (NISP)……………………………………………………………………………………..158

Figure 6.10: Element distribution of marine turtles expressed in number of identified specimens (NISP)…………………………………………………………………………160

Figure 6.11: Element distribution of marine turtles expressed in number of identified specimens (NISP) Macrounit 2b……………………………………………………...... 165

Figure 6.12: Element distribution of Iguanas expressed in number of identified specimens (NISP) Macrounit 2b ……………………………………………………………………..166

Figure 7.1: Fish skeleton nomenclature By Debbi Yee Cannon (1987) and Perdikaris et al (2004)……………………………………………………………………………………...175

Figure 7.2: Fragmentation size classes used for classification of bone state of preservation . 178

Figure 7.3: The cured fish model (photo Y. Krivogorskaya)……………………………….181

Figure 7.4: Fish filleting (Peebles et al 2010)...... 185

Figure 7.5: Fish butchered by method 2 (left- center) and method 1(right) (Zohar and Cooke 1997)……………………………………………………………………………………...186

Figure 7.6: Frequent genera 1/8‖ mesh (excluded genera < 1%)……………………..…....192

Figure 7.7: Frequent genera 1/16‖ mesh (excluded genera <1%) ...... 192

Figure 7.8: The relative abundance of the species in 1/8‖ mesh at Vampiros 1 and 2……..195

Figure 7.9: The relative abundance of the species in 1/16‖ mesh at Vampiros 1 and 2 ...... 196

Figure 7.10: The most frequent species from 1/8‖ mesh(%NISP excluding < 1%)...... 197

Figure 7.11: The most frequent species from 1/16‖ mesh (%NISP excluding < 1%)……..199

Figure 7.12: The relative abundance of the species (NISP) at Vampiros 1 and 2 by fish estimated biomass ...... 220

Figure 7.13: Localization of some natural and cultural bone modifications in Vampiros -1 and 2 stratigraphic profiles...... 223

Figure 7.14: Vampiros 1and 2. Preservation and modifications: a. deformed vertebra, b, c &d. different type of scales and e. branchial bones...... 226

Figure 7.15: Macrounit 2, Family Ariidae: a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)...... 231

Figure 7.16: Survival index (SI) for Ariidae fish remains from Macrounit 2...... 232

Figure 7.17: Macrounit 2, Family Carangidae: a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each) ...... 234

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Figure 7.18: Survival index (SI) for Carangidae fish remains from Macrounit 2...... 235

Figure 7.19: Macrounit 2, Family Clupeidae: a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)...... 236

Figure 7.20: Survival index (SI) for Clupeidae fish remains from Macrounit 2...... 237

Figure 7.21: Macrounit 2, Family Haemulidae a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)……………………………...………………... 238

Figure 7.22: Survival index (SI) for Haemulidae fish remains from Macrounit 2...... 239

Figure 7.23: Macrounit 2, Family Polynemidae a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)...... 240

Figure 7.24: Survival index (SI) for Polynemidae fish remains from Macrounit 2...... 241

Figure 7.25: Macrounit 2, Family Sciaenidae a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)……………………………………...... …….…242

Figure 7.26: Survival index (SI) for Sciaenidae fish remains from Macrounit 2...... 243

Figure 7.27: Macrounit 2, Family Pristigasteridae a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)………………………….……244

Figure 7.28: Survival index (SI) for Pristigasteridae fish remains from Macrounit 2...... 245

Figure 7.29: Macrounit 2, Family Belonidae: a. Element distribution, b. Comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. Vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)…………………………………………...... …246

Figure 7.30: Survival index (SI) for Belonidae fish remains from Macrounit 2...... 247

Figure 7.31: Macrounit 2, Family Tetraodontidae: a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)……………………………….248 xxii

Figure 7.32: Survival index (SI) for Tetraodontidae fish remains from Macrounit 2...... 249

Figure 7.33: Articulated bones: a. Ariidae Neurocranium, b. Clupeidae Caudal vertebrae,c. Ariidae ceratohyal-epihyal-hypohyal, (red arrow indicate cut mark)d.& e. Ariidae and Carangidae skull bones, f. Tetraodontidae Dentary-articular, g. Ariidae Quadrate-opercle, h. Ariidae vertebral complex, i. Ariidae Coracoid- cleithrum...... 251

Figure 7.34: Ariidae NISP vertebrae Macrounit 2= 5, Belonidae NISP vertebrae Macrounit 2= 6...... 255

Figure 7.35: Carangidae NISP vertebrae Macrounit 2= 398, Haemulidae NISP vertebrae Macrounit 2= 40...... 255

Figure 7.36: Polynemidae NISP vertebrae Macrounit 2= 80, Pristigasteridae NISP vertebrae Macrounit 2=15...... 256

Figure 7.37: Figure 7.34: Sciaenidae NISP vertebrae Macrounit 2= 15 Tetraodontidae NISP vertebrae Macrounit 2= 87…………………………………………256

Figure 7.38: Fine cut marks: a. HaemulidaeVertebra cut mark, b. Tetraodontidae Supracleithrum´s cut mark, c.& h. SciaenidaeVertebrae cut mark, d. & e. Ariidae Ceratohyal cutmark, f. Shark vertebrae cut mark, g. Sciaenidae Anal spine post-depositional cut mark...... 265

Figure 7.39: Chopped marks and fractures: a. Fractured dorsal spine (Ariidae), b. & d. transverse fractures in cleithrum (Ariidae), c. longitudinal fracture in ethmoid (Ariidae), e. transversal fracture in meptaterygoid (Ariidae), f. longitudinal fracture premaxilla (Tetraodontidae), g. Chopped coracoid (Ariidae), h. Chopped frontal (Tetraodontidae), i. Chopped supraoccipital (Ariidae)………………………………………………………….276

Figure 7.40: Ariidae specimens from Vampiros Macrounit 2. NISP= 835...... 278

Figure 7.41: Carangidae specimens from Vampiros Macrounit 2. NISP= 1065...... 279

Figure 7.42: Clupeidae specimens from Vampiros Macrounit 2. NISP= 84...... 280

Figure 7.43: Haemulidae specimens from Vampiros Macrounit 2. NISP=201...... 281

Figure 7.44: Polynemidae specimens from Vampiros Macrounit 2. NISP= 291...... 282

Figure 7.45: Sciaenidae specimens from Vampiros Macrounit 2. NISP= 320...... 283

Figure 7.46: Pristigasteridae specimens from Vampiros Macrounit 2. NISP= 140...... 284

Figure 7.47: Belonidae specimens from Vampiros Macrounit 2. NISP= 130...... 285

Figure 7.48: Tetraodontidae specimens from Vampiros Macrounit 2. NISP= 442...... 286

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Figure 8.1: Sitio Sierra, Family Clupeidae: a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)...... 295

Figure 8.2: Survival index (SI) for Clupeidae fish remains from Sitio Sierra...... 296

Figure 8.3: Sitio Sierra, Family Haemulidae: a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)...... 297

Figure 8.4: Survival index (SI) for Haemulidae fish remains from Sitio Sierra...... 298

Figure 8.5: Sitio Sierra, Family Carangidae: a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)...... 299

Figure 8.6: Survival index (SI) for Carangidae fish remains from Sitio Sierra...... 300

Figure 8.7: Sitio Sierra, Family Ariidae: a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)...... 301

Figure 8.8: Survival index (SI) for Ariidae fish remains from Sitio Sierra...... 302

Figure 8.9: Sitio Sierra, Family Sciaenidae: a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)...... 303

Figure 8.10: Survival index (SI) for Sciaenidae fish remains from Sitio Sierra...... 304

Figure 8.11: Ariidae NISP vertebrae = 7, Carangidae NISP vertebrae=734………………..305

Figure 8.12: Clupeidae NISP vertebrae = 74, Haemulidae NISP vertebrae =744...... 306

Figure 8.13: Figure 8.13: Sciaenidae=24…………………………………………………...306

Figure 8.14: AG-125, Family Ariidae: a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)...... 312

Figure 8.15: Survival index (SI) for Ariidae fish remains from Ag-125...... 313

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Figure 8.16: AG-125, Family Sciaenidae: a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)...... 314

Figure 8.17: Survival index (SI) for Sciaenidae fish remains from Ag-125...... 315

Figure 8.18: AG-125, Family Carangidae: a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)...... 316

Figure 8.19: Survival index (SI) for Carangidae fish remains from Ag- 125………………………………………………………………………………………...317

Figure 8.20: AG-125, Family Haemulidae: a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)...... 318

Figure 8.21: Survival index (SI) for Haemulidae fish remains from Ag-125...... 319

Figure 8.22: AG-125, Family Tetraodontidae: a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)...... 320

Figure 8.23: Survival index (SI) for Tetraodontidae fish remains from Ag-125...... 321

Figure 8.24: AG-125, Family Clupeidae: a. element distribution, b. comparison of relative proportions of the cranial and axial bones recovered. In a whole fish, the relative proportions would be equal, c. vertebral series. A complete fish skeleton would produce a graph of exactly equal proportions for % MAU (33% each)...... 322

Figure 8.25: Survival index (SI) for Clupeidae fish remains from Ag-125...... 323

Figure 8.26: Carangidae NISP vertebrae = 27, Haemulidae NISP vertebrae=13……….….324

Figure 8.27: Sciaenidae NISP vertebrae = 34...... 324

Figure 8.28: a. cut mark on Ariidae´s caudal vertebrae, b. Ariidae´s cleithrum showing chopped and impact marks and, c. Sciaenidae´s premaxilla chopped...... 327

Figure 8.29: Ariidae specimens from Sitio Sierra. NISP= 2535...... 346

Figure 8.30: Carangidae specimens from Sitio Sierra. NISP= 2229...... 347

Figure 8.31: Clupeidae specimens from Sitio Sierra. NISP= 3824...... 348 xxv

Figure 8.32: Sciaenidae specimens from Sitio Sierra. NISP= 158...... 349

Figure 8.33: Haemulidae specimens from Sitio Sierra. NISP= 3619...... 350

Figure 8.34: Ariidae specimens from Ag-125. NISP= 232...... 351

Figure 8.35: Carangidae specimens from Ag-125. NISP= 77...... 352

Figure 8.36: Clupeidae specimens from Ag-125. NISP= 87...... 353

Figure 8.37: Haemulidae specimens from Ag-125. NISP= 60...... 353

Figure 8.38: Sciaenidae specimens from Ag-125. NISP= 34...... 354

Figure 8.39: Tetraodontidae specimens from Ag-125. NISP= 63………………………….355

Figure 9.1: Vampiros-2: hearth surrounded by postholes (toothpicks) by Pearson………...373

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List of Symbols, Abbreviations and Nomenclature

Symbol Definition U of C University of Calgary PSM The Proyecto Santa Maria ITCZ Inter-tropical Convergence Zone ENSO El Niño Southern Oscillation STRI Smithsonian Tropical Research Institute NISP Number of Identified Specimens MNI Minimal Number of Individuals MAU Minimal Animal Units SI Survival Index WMI Weighted mean index of fragmentation ETH Ethmoid (Lateral ethmoid) PRF Prefrontal VOM Vomer SET Supraethmoid ALI Alisphenoid PAR Parasphenoid SPH Sphenotic PTE Pterotic EPI Epiotic OPI Opistothic PRT Prootic OTO Otolith PAI Parietal NAS Nasal FRN Frontal SCALE Scale bone SPB Orbitals LAC Lachrymal DEN Dentary ANG Articular RET Retroarticular PRO Preopercle OPE Opercle SBO Subopercle INT Interopercle BR Branchiostegal ray PAL Palatine ECT Ectopterygoid EPU Epural EHS Expanded haemal spine PEN Penultimate vertebrae SUP Supraoccipital xxvii

EXO Exoccipital MTR Mesopterygoid MET Metapterygoid HYO Hyomandibular SYM Sympletic INH Interhyal EPH Epihyal CER Ceratohyal HYH Hypohyal BAH Basihyal PP Pharyngeal plate EPB Epibranchial CEB Ceratobtanchial HYP Hypobranchial BAB Basibranchial URO Urohyal PHA Pharingobranchial. POS Postemporal SPC Supracleithrum CLE Cleithrum PCM Postcleithrum QUA Quadrate RAD Radials BAM Basipterygium IS Interhaemal spine PRECAUDAL Precaudal vert. ULTIMATE Ultimate vert. HRP Hypural ENS Expanded nrl. spine BAS Basioccipital COR Coracoid VX Vertebral complex DE Dorsal spine PETI Dorsal pterygiophore PTG Pterygiophore EP Pectoral spine SKULL Unidentified skull frag. SPINE Unidentified spine frag. S Scapula SS Spiny scale PM Premaxilla GLOSS Glossohyal ENT Entopterygoid BPH Basisphenoid

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Appendices

Appendix 1: Ariidae skeletal elements in a complete fish...... 430

Appendix 2: Haemulidae skeletal elements in a complete fish...... 432

Appendix 3: Carangidae skeletal elements in a complete fish...... 434

Appendix 4: Polynemidae skeletal elements in a complete fish...... 436

Appendix 5: Sciaenidae skeletal elements in a complete fish...... 438

Appendix 6: Clupeidae skeletal elements in a complete fish...... 440

Appendix 7: Tetraodontidae skeletal elements in a complete fish...... 442

Appendix 8: Belonidae skeletal elements in a complete fish...... 444

Appendix 9: Pristigasteridae skeletal elements in a complete fish...... 446

Appendix 10: (on CD) Primary data

Appendix 11: (on CD) Secondary data

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Chapter 1

Introduction

My thesis is an attempt to determine the function of a site, Cueva de Los Vampiros, situated along the central Pacific coast of Panama and to understand its role in the regional economy.

Based on my research at the site and analysis of the faunal remains, I will argue that the site existed principally for the purpose of processing fish for preservation and possibly for transport to sites further inland. Although Archaeological and ethnohistorical records indicate that fish and fish products were important exchange commodities within chiefdoms at the moment of the Spanish contact (Cooke, et al. 2008), the occupation at Cueva de Los

Vampiros pre-dates the evidence for the rise of chiefdoms in Panama by several centuries.

My thesis, then, aims to probe the antiquity of the technology and practices that presaged the trade and redistribution of fish in the economies of the subsequent Panamanian chiefdoms.

The exchange of foodstuffs is one aspect of trade that has received less attention than it merits.

The survival of fish remains in archaeological contexts is rare in Panama; the exception is the central Pacific coast of Panama where several archaeological sites ranging in age from 7000 to 4500 years BP have fish remains in domestic and mortuary contexts and

Chiriquí sites such as Cerro Brujo, Sitio Drago and La Pitahaya (Cooke, et al. 2008; Cooke and Ranere 1992a; Jiménez and Cooke 2001; Linares and Ranere 1980; Wake et al 2004). In

2004 I participated in excavations at two rockshelters at the site of Cueva de los Vampiros

(Figure 1.1), - near the current coastline of Parita Bay. Occupations at these shelters range back to Paleoindian times, the excavation of which was the main objective of Dr. Georges

Pearson, whom I was assisting (Cooke and Ranere 1984; Pearson, et al. 2003). Stratified deposits in both shelters above the Paleoindian component contained Ceramic period remains with abundant and well-preserved fish remains. The analysis and interpretation of these remains constitutes the major part of this thesis.

Figure 1.1: Map of the Study area (from Google maps)

Coastal Resources

Much of the research done in environmental archaeology has seen coastal resources such as fish, shellfish, marine mammals and birds as vital elements for the dietary welfare of Pre-

Columbian cultures in the Americas. As Pre-Columbian inhabitants began to produce food 2

crops, plants rich in carbohydrates became dietary complements to the protein-rich sea foods (Gupta 2004; Hayden 1992; Kabo 1985; Richerson 2001). There were only a few domesticated animals in the Americas: camelids (Lama glama, Vicugna pacos) and guinea pig

(Cavia porcellata) in Andean South America; turkey (Meleagris pavo) in Mesoamerica and

Southwesta, dog (Canis lupus familiaris) were introduced from Asia, and Muscovy duck (Carina moschata) which has a wide distribution in both continents (Cooke, et al. 2008; Emery 2004;

Keegan 1989; Leonard, et al. 2002; Mengoni Goñalons 1999, 2004; Pendergast 2004; Stahl

2003; Stahl 2005). Humans, then, relied mainly on fishing and hunting for protein.

The particular problem I have chosen to address within the larger debates about the relative merits of marine resources in human diets concerns the potential of estuaries to provide aquatic resources for people who live too far from the coast to be able to exploit estuarine resources directly. Ecosystems such as cool upwelling coastal waters and the tropical coasts and estuaries provide local populations with abundant and easily obtained protein, as well as fat (Leach 2006a). According to archaeological research in the Americas, estuaries were focal points for early sedentism and population growth, and they retained their advantages for human settlement even after extensive agriculture based on maize and root crops became the primary subsistence activity (Bryan 1993; Drennan 1996; Stahl and

Oyuela Caycedo 2007; Stothert 1985).

Within the scholarly divide of opinion on the importance of aquatic resources, my research places me with those that Erlandson (2001) calls the "Garden of Eden" theorists, whose intellectual precursor is Carl Sauer (1966). These scholars praised the abundance of food and ease of capture of prey in aquatic habitats, and related these factors causally to the establishment of large and sedentary villages, the development of food production, and the 3

emergence of chiefdoms as related to these factors (Bearez 1996; Carneiro 1995; Oyuela-

Caycedo 1996; Oyuela-Caycedo and Bonzani 2005). Other scholars take an opposite position which is called the "gates of Hell" approach by Erlandson. Conversely, this approach developed initially by Osborn (1977) relegates shellfish and other aquatic foods to the status of "marginal" resources that are irregularly distributed, expensive to harvest or process and also nutritionally poor in comparison to terrestrial game. Aquatic resources are seen as desparate measure.

The Pre-Columbian exploitation of shellfish, fish, birds and mammals on the coasts and tropical Pacific estuaries goes back to the Holocene and further in Chile, Peru, Ecuador,

Colombia, Panama, Central America and the Antilles, and continued until the Spanish

Conquest. Also the earliest settlements near estuaries like Las Vegas, Cerro Mangote,

Monagrillo, Chantuto and Puerto Hormiga, provide evidence for plant utilization (Bearez

1996; Byrd 1976; Cardera, et al. 2007; Cooke and Jiménez 2004; Cooke, et al. 2008; Cooke and Ranere 1999; Cooke and Sanchez Herrera 2004; Cooke 1992, 2004a, b; Davis 1982; deFrance 1989; Dillehay 2009; Grouard 1995; Keegan 1989; Legros 1992; Lippold 1991;

Moravetz 2003; Peres 2001; Petersen 1997; Reitz and Masucci 2004; Rewniack 2006; Scudder

1995; Stahl 2003; Stahl and Oyuela Caycedo 2007; Stothert 1985; Sutty 1995; Voorhies 2004;

Watters 1989; Wing 1995, 2001).

The economic importance of coastal resources has been stressed particularly in

Peruvian archaeology, where evidence for marine fishing in a prolific coastal environment goes back to 11,000 radiocarbon years ago (rcya) (deFrance 2009, 2001; Moseley 1992a, b;

Sandweiss, Keefer, et al. 1999; Sandweiss 1996; Sandweiss, et al. 1989; Sandweiss, et al.

1998). In agreeing with the ―Garden of Eden‖ theorists, I believe that protein and fats 4

provided by aquatic animals were fundamental complements to the largely plant-based diets of Pre-Columbian societies in the Americas. This conclusion has been supported by decades of research into the topic, while Osborn's work has been shown to be ecologically naive as he concentrated almost exclusively on shellfish and ignored the huge potential for aquatic resources with terrestrial animals and plant resources. Aquatic resources whose productivity, niche diversity availabilty all year round and ease-of-exploitation are considered by many scholars to have been important for the development of early sedentary societies (deFrance

2009, 2001; Feldman 1992; Marcus, et al. 1999; Quilter 1991; Reitz 1988, 2001; Reitz and

Masucci 2004; Sandweiss, et al. 1999; Sandweiss et al. 1999; Sandweiss 1996; Sandweiss, et al.

1989; Sandweiss, et al. 1998).

Marine resources, in conjunction with plants and terrestrial animals, have played a vital role in the social and demographic processes that fostered social complexity in the

Intermediate Area like the small scale Panamanian societies (Capriles, et al. 2008; deFrance

2009; Drennan 1991; Drennan 1996; Hoopes 2004; Raymond 2008). I (and other scholars) acknowledge that the relationship between aquatic environments and human cultural and economic development is far more complex than a simplistic ‗Garden of Eden‘ scenario indicated by many of the references cited above. Pre-Columbian inhabitants, who lived inland, obtained aquatic resources from many sources. Along the major river systems of tropical South America like the Amazon Basin, freshwater fish faunas are diverse and contain taxa that attain very large sizes, providing a considerable dietary contribution

(Chernela 1993; Jackson 1983; Peña León 2007; Smith 1981; Vieco and Oyuela-Caycedo

1999). Some Central and Southamerican lakes, e.g., Lake Nicaragua, and South American lakes, e.g., Lake Titicaca, supported productive fisheries (Capriles, et al. 2008; Healy and Pohl

1980; Peña León 2007; Rewniack 2006; Villa 1982). Freshwater rivers, however, are much less productive for example on the Central American isthmus, especially on the Pacific side where rivers lack large amphidromous species that are present in Caribbean basins (e.g.,

Joturus, Megalops) (Cooke and Jiménez 2008; Cooke, et al. 2008; Cooke and Ranere 1992a;

Cooke and Ranere 1999; Cooke and Tapia-Rodríguez 1994; Villa 1982).

Fish and Trade

The Pacific coast of Panama offers a potentially informative laboratory for examining the beginnings of trade in fish and fish products from the source to peoples whose locations make direct access problematic or impossible. An efficient way of compensating for the lack of aquatic protein sources in inland locations is to preserve and transport the abundant inshore marine fish. Drying, salting and smoking are the most effective means of preservation.

My experience working at the Vampiros shelters on the coast of Parita Bay alerted me to the possibility of applying zooarchaeological techniques to the question of when and how large quantities of estuarine fish resources may have been moved inland. A prior analysis of part of a stratum at Vampiros-1 by Cooke indicated that nearly all the vertebrate materials belonged to fish (Cooke 1988; Cooke, et al. 2008). Although it is of course possible that these remains represent food waste discarded by people who went to this shelter to undertake other activities, certain details, i.e., burnt bones and lots of whole and articulated

(fused) bones, suggested that this site's function may have been to smoke and/or salt fish for subsequent transport inland, either by the shelter's occupants themselves or through some kind of exchange mechanism. For comparison, I chose to study the fish remains at the site

of Sitio Sierra, a village-size settlement contemporary with the Vampiros shelters and situated a 13 kilometres ways inland. There are not contemporary sites further inland contemporaneous with these sites.

The transport of animal body parts has been studied by archaeologists in order to understand the flux between producer and consumer sites; however, much of the literature focuses on either large or medium-sized vertebrates, mostly mammals or molluscs.

Inferences are based on skeletal element distributions recovered from archaeological sites.

The recovered elements provide insight as to site function, and methods of food preparation, to name a couple (Bartram and Marean 1999; Binford 1977, 1978, 1981, 1984;

Binford, et al. 1983; Bird and Bird 2000; Klein, et al. 1999; Lupo 2001; O'Connell, et al.

1990; Reitz and Massucci 2004; Waselkov 1987; Whitridge 2001; Wing 2001). However, there has been less discussion regarding the transport or trade of fish. The published research concerning fish has focused on procurement strategies, subsistence, catchment areas and fishing technology (Ceron-Carrasco 2005; deFrance 1989; Fernández 2004;

Gutierrez Gonzalez 1997; Kerbis 1980; Leach 2006c; Tourunen 2008; Voorhies 2004; Wing

1995, 2001; Wing and Wing 1995). However, indirect evidence of the transport of fish has been suggested by the presence of non- local taxa (Bearez 1996; Masucci 1995; Quilter

1991; Reitz 2001; Sandweiss, et al. 1999; Sandweiss, et al. 1989; Sandweiss, et al. 1998;

Wheeler and Jones 1989), and by more direct evidence such as differential distribution of skeletal elements (Belcher 1998; Carr 1986, 1987; Cooke 1988; Gotz 2008; Tourunen 2008;

Zohar and Cooke 1997; Zohar, et al. 2001), the sizes of the fish represented (Barrett 1997), the presence of cut marks (Willis, et al. 2008) and complementary archaeological evidence

(i.e artefacts)(Bearez 1996; Leach 2006b; Masucci 1995; Reitz and Masucci 2004). 7

Recently, several zooarchaeologists in Norway and Korea have been studying the use of fish remains, mainly butchering practices and exchange, in order to explain food practices, ethnicity and trade (Amundsen 2008; Barrett, et al. 2008; Krivogorskaya, et al. 2005;

Krivogorskaya., et al. 2005; Norton, et al. 1999; Perdikaris 1999).

Many of the above studies have looked for analogues in the recent or contemporary ethnographic record with which to compare and contrast zooarchaeological data (Acheson

1981; Alegret Tejero 1989; Belcher 1998; Bird and Bird 2000; Gragson 1992; Knudson, et al.

2004; Lieber 1994; Smith 1981; Stewart and Gifford-Gonzalez 1994; Zohar and Cooke

1997). As individual activities, fishing and preparing fish products are considered primarily a household, day-to-day activity. But fishing may be included with other subsistence activities requiring specialization and thus interdependence of household members or households

(Cooke 1998, 2001; Lieber 1994; Moss 1993; Perdikaris 1999; Reitz and Massucci 2004;

Titcomb 1972; Waselkov 1987). Several examples in the anthropological literature suggest that individual households have control of fishing territories which supply their own communities (Acheson 1981; Durrenberger and Pálsson 1987; Jackson 1983; Levieil and

Orlove 1990; Lieber 1994; Vieco and Oyuela-Caycedo 1999). Ethnohistoric documents suggest that elites in Pre-Columbian Panama had special hunting grounds ("cotos"). It is likely that fishing grounds were also subjected to social controls (Cooke 1998; Jopling 1993).

Anthropological studies mention several models, regarding how goods like raw materials or manufactured goods (pottery and stone tools), prestige items and foodstuff (fish and salt) were exchange depending of the complexity of the socio-political institutions, distance (internal versus long distance) and other factors. Sahlins (1972), for example 8

mentions a model of commercial movement depending of social relationships: ―the generealized reciprocity‖ is between people of the same household, ―the balanced reciprocity‖ is between people of the same tribe and ―the negative reciprocity‖ is between people of different tribes. Polanyi (1957) refers to exchange activies associated with social relationships such as reciprocity with egalitarian social relations, redistribution where there is political centralization and direct commerce with is related to the modern world.

Finally, Renfrew (1975) based on spatial relationships , settlements and material findings suggests three models of internal trade: ―the direct access‖ where people collect the resources ,‖ the home base reciprocity‖ where people from one area visit people from another area, and exchange items in which ―the boundary reciprocity‖ implies that two groups meet at a common limit and exchange items. His five models of external trade are the following: ― the down-the line exchange‖ where each village of the line would receive the goods through exchange from the nearest neibourg to the source, ― the central place redistribution and the central place market exchange‖ where groups give items to a central authority or market respectively, ―the prestige-chain exchange‖ where the distribution of prestige artefacts are transferred to others in a continuous process as offerings in a burial, ― the freelance commercial trade‖ which involves long distance and independent entrepreneurs who transferred objects that are not high prestige and ― the directional commercial trade‖ where the commodities are transferred from a specific source to a specific destination.

Ethnohistorical and archaeological accounts in the Americas mention mechanisms of exchange for gold, obsidian tools, fine ceramic vessels, jade, feathers, salt, textiles, chocolate, 9

thorn oyster and fish. This literature also mentions specialied merchants (pochtecas, Siguas and mindales) who travel by land or sea, markets and tribute payments (Coe and Koontz

2002; Cooke and Sanchez Herrera 2001; Demarest 2004; Earle 2010; Lothrop 1942b;

Marcos 1986; Martin 2001; McEwan and Delgado Espinoza 2008; Smith 1990). However as

Olga Linares (1977:71) points out the archaeological evidence of internal trade in food such as fish is hard to determine since this kind of exchange leaves few traces, in contrast to the writing evidence which describes this activity at the time of the conquest (see chapter three).

Objectives

In Panama vertebrate bones obtained in 2005 at the coastal site Cueva de los Vampiros were taxonomically, anatomically, and ontogenetically identified from two localities. Additional assemblages from the inland site Sitio Sierra, noted above, and a late coastal site AG-125 were also studied. In this dissertation I ask whether Vampiros deposits result from processing fish for export. I focus on results from Vampiros, but I compare them with Sitio

Sierra, and AG-125. This comparison illuminates distinctive aspects of the Vampiros deposit and facilitates a better understanding of taphonomic biases.

I focus my attention on those aspects of the distribution of fish taxa and body parts, which point toward the preparation of fish for preservation in terms of what might have been traded, and I address the following specific questions with zooarchaeological material from Vampiros:

1) Is there evidence for the preservation of fish, i.e., smoking and drying (e.g., physical damage, differential exposure to heat, and differential body part distribution)? 10

2) Are their changes through time in fish abundance suggestive of corresponding changes in habitat use and/or fishing techniques?

3) What was the role of the Vampiros shelters in the local and regional subsistence economy?

This is a research project in tropical America that focuses on the relationship between fish diversity, taphonomy and human behavior in Pre-Columbian times. It builds on ethnoarchaeological studies conducted by Irit Zohar and Richard Cooke (1997) at nearby modern coastal fishing villages. Their work provided empirical data on the relationship between fish preparation techniques, fish size, bone loss and damage, and bone

"survivability." My project will add a new behavioral dimension to our knowledge of Pre-

Columbian human adaptations around Parita Bay, where so much research has been conducted since the 1940s ( Carvajal Contreras 1998; Carvajal Contreras, et al. 2008; Carvajal

Contreras, et al. 2005; Carvajal Contreras and Hansell 2008; Cooke 1977, 1989; Cooke 1993;

Cooke, et al. 2001; Cooke, Isaza, et al. 2003; Cooke and Jiménez 2004; Cooke, et al. 2000;

Cooke, et al. 2007; Cooke, et al. 2008; Cooke and Ranere 1984; Cooke and Ranere 1989;

Cooke and Ranere 1992a, b, c; Cooke and Ranere 1999; Cooke, Sanchez, et al. 2003; Cooke, et al. 2000 ; Díaz 1999; Dickau 2005; Dickau, et al. 2007; Haller 2004; Hansell 1979, 1988;

Ichon 1980; Isaza-Aizpurua 1993; Isaza-Aizpurua 2007; Jiménez 1999; Jiménez and Cooke

2001; Ladd 1964; Lothrop 1937, 1942a ; Lyall 2007; McGimsey 1956; Mayo and Cooke 2005;

Peres 2001; Ranere and Cooke 1991; Ranere and Cooke 1996; Ranere and Cooke 2003;

Sanchez Herrera 1995; Sanchez Herrera and Cooke 2000; Willey, et al. 1954). Because of the complementary research that has been carried out in this area over the past few decades, my research, I hope will provide a basis for understanding the origins of fish harvesting and 11

preservation that later resulted in the widespread transport and redistribution of marine resources among inland communities and within the broader context of Pre-Colombian tropical chiefdoms.

Chapter 2 is a general presentation of the study zone, Cueva de los Vampiros, in regard to past and present environments as well the history of research conducted at this site. This chapter also describes provenience, methods, and techniques used during the excavation of both shelters.

The third chapter presents the temporal and cultural contextualization of Cueva de los Vampiros. It is a general overview of Central Pacific Panama prehistory and provides ethnohistorical evidence of the exchange of commodities such as fish.

The fourth chapter addresses the cultural remains recovered from Cueva de los

Vampiros and seeks to correlate the lithic and pottery typologies with the radiocarbon dates, as well as provides indirect evidence of site activities, function and formation process.

In the fifth chapter the invertebrate remains and methods of analysis are described, and interpretations of subsistence, preferred habitats and procurement techniques are presented.

In the sixth chapter, analytical methods for describing non-fish vertebrate remains as well as the interpretation of subsistence, habitats and taphonomy are presented.

Vampiros fish remains are described in chapter seven. To test the possibility that

Vampiros was a site for processing and curing fish, the analysis focused on the distribution

of fish taxa, body parts, physical damage to bones, differential exposure to heat and distribution of cut marks.

To provide comparative data to investigate the exchange and production of cured fish at Cueva de los Vampiros, chapter eight describes Sitio Sierra and AG-125 fish assemblages. AG-125 may have replaced the Vampiros shelters as the primary fishing settlement at the mouth of the Santa María River and may present similar butchering patterns. Sitio Sierra, on the other hand, overlaps in time with the later stages of the fishing camp at Vampiros and may indicate a habitation site where Vampiros production was distributed.

Finally, chapter nine provides a synthesis and conclusions of the preceding chapters as a foundation for an evaluation of the evidence for preservation and transport of fish, changes in habitat, and the role of the Vampiros shelters in the local and regional economy.

Chapter 2

Cueva de los Vampiros Site (AG-145)

Introduction

Vampiros is an archaeological site which consists of several rockshelters. This research covers two localities: Vampiros 1- and Vampiros- 2. Both are situated near Parita Bay on the central Pacific coast of the Isthmus of Panama (Figure 2.1). Today, they are located on the

Santa Maria estuary on a small rocky outcrop or inselberg called Cerro Tigre some 2.6 kilometers from the present-day shoreline and standing 0.3 kilometers in straight line from the north bank of the Santa Maria River. These sites are located in the Eastern Pacific biogeografic region.

Figure 2.1: Map and aerial photography showing the location of the Vampiros-1 and Vampiros-2 shelters with respect to other Pre-Columbian sites around Parita Bay. AS - Aguadulce Shelter, CJD - Cerro Juan Diaz, CM - Cerro Mangote, LA - Cueva de los Ladrones, LS-10 - Las Huertas, LS-15 - La Isleta, LA-29 - La Chilonga, MO - Monagrillo, SS - Sitio Sierra, VA - Vampiros shelters, ZA – Zapotal (Carvajal et al 2008).

Modern Environmental Setting

The Pacific coast on the Parita Bay exemplifies a typical Central American coast characterized by biological and hydrological heterogeneity. Currently, the Vampiros rockshelters are situated in an estuarine ecosystem. Parita Bay is c-shaped, resulting from a marine invasion of the Pacific continental shelf which at this area is approximately 25 km wide (Figure 2.2). This Eastern Pacific biogeographic region of Panama is highly variable, rich in open-sea fish species but has a poor development of coral reefs (Allen and Robertson

1998; Barber 1981; Jackson 1997). However, the estuarine ecosystem near Vampiros is characterized by discharges of fluvial sediments, broad tidal ranges that form long muddy zones with a patchy distribution of habitats. As a result of its rich natural structure, this ecosystem houses a diversity of species of fish, molluscs, birds, reptiles, mammals and plants

(Cooke 2002; D´Croz and O‘Dea 2007).

One of the elements of this ecosystem is the Santa Maria River. It is a 145 km long perennial river; its watershed covers approximately 3315 km2. This river runs west to east along a wide coastal plain, characterized by changing oxbows and seasonal freshwater swamps, as well as remnants of dry forest, gallery forest, and Avicennia mangroves. During the dry season, which runs from December to April, the alvina (or salt marsh) of El Tigre is exposed to aeolian erosion. The Santa Maria River discharges into a delta in Parita bay that has an average of 1300mm of rainfall during the rainy season (May- November). The weather is seasonal, characterized by wet-dry tropical (Aw) climate with an average annual temperature range from 23-27 °C (Barber 1981; Palka 2005).

The bay as I mentioned above is located in the Eastern Tropical Pacific region. This biogeographic region is characterized by a complicated interaction between seasonal atmospheric and physical oceanographic conditions which influences the fisheries of the area. First, trade winds converging into the ITCZ (Inter-tropical Convergence Zone) are responsible for the rainfall regime producing the wet and dry seasons. These seasonal winds interact with upwellings which occur during the dry season (December to April). These upwellings are cool and nutrient rich water. The seasonal upwelling is stronger in the Pearl

Islands than in Parita Bay. El Niño Southern Oscillation (ENSO) also interacts with the trade winds. ENSO is characterized by an increase in water temperature that alters the temperature and salinity of the estuary and has an enormous impact in the area every three or eight years.

In the study area many marine currents interact which explains the heterogeneity and richness of the Eastern Tropical Pacific. Two west horizontal movements of seawater at the ocean's surface, the South Equatorial Current (SEC) and the Northern/Southern Subsurface

Counter Current are associated with a recirculating tropical gyre. These currents modified seasonally the salinity and temperature of the ocean (Allen and Robertson 1998; Kessler

2006 see Figure 5:187; MacIlvaine and Ross 1973: 215).

The upwelling, currents and atmospheric events change the productivity of the sea

(e.g., nutrients, plankton and pelagic fishes) (Bemis and Geary 1996; D´Croz, et al. 1991;

D´Croz and O‘Dea 2007; D´Croz and Robertson 1997; Jackson and D'Croz 1997).

Nowadays, Parita Bay is a dynamic landscape as well as in the past. Parita Bay coastal geomorphology has changed since the early Holocene, influenced by the low relief of the deltas, the constrained river channels, tectonic uplift of the upper costal plain, sedimentation rates of the bay, sea level fluctuations, climatic change, and anthropogenic activity (Barber

1981; Dere 1981).

The low-energy coastline of Parita bay has a tidal range average of 4.5 m most of the year, but which can range up to 6m (locally called aguajes). Extended intertidal mudflats, approximately 3 km wide, produce complex and uneven geomorphologic zones (supratidal, high-tidal, mid-tidal and low-tidal) favouring not only mangrove development but diverse micro-environments and their component fish species (Barber 1981; Carvajal Contreras, et al. 2008; Clary, et al. 1984; Cooke and Ranere 1999; Cooke 1992a).

Figure 2.2: Map of the study area showing environmental setting surrounding Vampiros rockshelters (AG-145), Sitio Sierra (AG-308) and

AG-125 (Instituto Tommy Guardia).

Paleo-environmental Setting

The young landscape of mudflats and mangroves that surrounds the Vampiros shelters rests on older geological formations. The geological basement is composed of Tertiary-age volcanic rocks such as lavas and tuffs. These extrusive rocks form inselbergs or monadnocks which are interbedded with Holocene marine and terrestrial sediments composed mainly of clay and sand. Among these inselbergs is Cerro El Tigre, the location of the Vampiros shelters (Barber 1981; Guardia 1988; Harmon 2005; Terry 1956).

The dynamic landscape of Parita Bay, characterized by changes in sedimentation and sea level, was studied by geologists during the Proyecto Santa Maria (PSM), a multidisciplinary project that analyzed prehistoric adaptations in this watershed. According to these scholars and other paleoecological evidence, the shoreline of Parita Bay is advancing into the Gulf of Panama as a result of sedimentation at the mouth of the Santa Maria River at a rate of 3 mm/yr-(Barber 1981; Clary, et al. 1984; Guardia 1988).

There are no indications of a human presence at Vampiros in sediments deposited between

15 190 ± 60 B.P. (16640[16210] 15820 cal B.C.) and a thin occupation floor with a bulk sediment date of 11550 ± 140 B.P. (12080–11980 [11520] 11950–11200 cal B.C.). Over this floor and beneath a charcoal date of 8970 ± 40 B.P. (8260 [8230] 8200 cal B.C.) Pearson

(2002) found Clovis overshoot flakes, a spurred end scraper, a thumbnailscraper, and a fluted bifacial point blade similar to South American fishtail fluted

Points. Cerro El Tigre and the Vampiros shelters were inland sites located approximately 50 km from the coast line (Figure 2.3). Pollen, phytoliths, clay and carbon records from La

Yeguada, Monte Oscuro and El Valle suggest that the cooler and drier Late Pelistocene climate, favored patchy and heterogeneous vegetation consisting of lowland woody scrub flora, savanna plants and gallery forests. Sea level stabilized during the Younger Dryas

(12500 to 11500 cal BP), during which time the sea was 50m below present levels. Simulated data suggests that rapid and small ENSO events occurred during this time (Bartlett and

Barghoorn 1973; Bush 2007). This environment was probably slightly modified by Clovis hunters. There is a modest evidence of burning activities at La Yeguada and there is no evidence at this time depth at Monte Oscuro (Piperno 2006; Piperno and Jones 2003).

Figure 2.3: Map of Panama´s coastlines during the Pleistocene (Instituto Tommy Guardia)

After these events and up to 8500 BP, the climate and vegetation is characterized by highest sea levels and moisture conditions. The sea level rose to 30 meters up the present 21

level (Barber 1981; Bartlett and Barghoorn 1973; Clary, et al. 1984; Lambeck, et al. 2002).

At this time, ENSO events became larger and less regular, continuing throughout the rest of the Holocene. As a result, there were more sustained wind patterns and a steadier equatorial upwelling regime (Bush 2007; González et al 2006; Koutavas, et al. 2006). Around 7000

BP, La Yeguada‘s paleoecological record indicates drier climatic conditions prevailed in

Central Panama, contemporary with a decline in secondary woody taxa and continued high levels of carbon. These changes are associated with the development of small scale cultivation of gardens around 9000 and 7000 B.P. in three Panamanian sites Aguadulce

Shelter, Carabali and Vampiros. Squash (Cucurbita), arrowroot (Maranta arundinacea), bottlegourd (Lagenaria siceria) and leren (Calathea allouia) phytoliths and starches were found in these Central Panamanian sites on stone tools. People intensified the slash and burn cultivation of these crops until 4000 BP (Bush 2007; Cooke 2995; Dickau 205; Dickau, et al.

2007; Piperno and Holst 1998; Piperno 2006; Piperno in press; Piperno, et al. 1992; Piperno and Jones 2003).

Dolores Piperno (1988:168–176; Piperno et al. 1985) identified pollen and phytoliths of domesticated maize (Zea mays) at Los Ladrones shelter dating to approximately 6000 B.C.

She also identified starch grain of domesticated manioc, and arrowroot, on edge-ground cobbles at Aguadulce Shelter dating between ca 6000–5000 B.C. (Piperno and Holst 1998:

771; Piperno and Pearsall 1998:219).

By 7000 yr BP, to a final period (5600 yr cal BP–Present) when sea level reached its present Parita Bay height, humidity persisted, and indicators of disturbance which is related to a global stabilization of the sea level, the ancient coastline would have been about 1.2 km

east of Cerro Mangote and ca. 4 km west of Cerro El Tigre (Figure 2.4). Coincidently at this time, nearby shell middens attest to marine adaptations on the central Pacific coast, e.g.,

Cerro Mangote(14C to cal. 5930–5450 B.C), Zapotal (cal. 1660–1220 B.C. (Beta-20850,

Cooke 1995: 173) and Monagrillo(cal 2880 [2610] 2460 B.C.), but Vampiros does not have cultural materials dating to this period; rather its stratigraphic sequence presents a hiatus for this period. Previous excavations at Vampiros suggest that this hiatus is associated with macrounit 3 (see stratigraphy below) when Cerro El Tigre was surrounded by the sea

(Gonzalez et al 2006; Cooke and Ranere 1999; Oyuela-Caycedo 1996; Pearson and Cooke

2002; Pearson 2002; Pearson, et al. 2003).

Figure 2.4: Hypothetic Parita Bay´s ancient coastline during 7000 BP

The Parita Bay deltas have been advancing seaward since 4000 BP faster at the Santa

Maria River (centre) than the Grande and La Villa rivers (edges). Between 2200 and 1900

BP, the sedimentation rate continued 1mm /yr at the Santa Maria delta. Information from the Gatun Basin indicates a rapid rise of the sea level around 2100 BP, followed by a recession between 2000 to 1000 BP (Bartlett and Barghoorn 1973). The landscape surrounding Parita Bay was transformed into anthropogenic savannas, heavily affected by agricultural activities. By this time (Figure 2.5), Vampiros would have been reoccupied and located near the active shoreline, which allowed early inhabitants to exploit sub-tidal and inter-tidal habitats (Barber 1981; Carvajal and Hansell 2008; Carvajal Contreras, et al 2008;

Clary, et al. 1984; Cooke and Ranere 1992, 1999; Golik 1968; Lambeck, et al. 2002; Nores

2004; Oyuela-Caycedo 1996; Suguio 1993; Weiland 1984). According to paleoenvironmental records, the climate was wetter and had more ENSO intervals from

180 B.C. to 550 A.D (Lachniet, et al. 2004; Piperno 2006).

Figure 2.5: Hypothetic Parita Bay´s ancient coastline during 2200 BP

The albina or salt flats appeared ca 1600 BP, and evidence of human activity decreased in the Vampiros rockshelters. After ca 2200 BP the archaeological record indicates a resettlement of inhabitants along the alluvial valleys (Figure 2.6). Vampiros was abandoned or less frequently occupied at this time, perhaps as a result of the increased distance from the active marine shore, which favored the establishment of new fishing stations such as AG-

125(Barber 1981; Carvajal Contreras, et al. in press; Clary, et al. 1984; Cooke and Ranere

1992c; Cooke and Ranere 1999; Golik 1968; Lambeck, et al. 2002; Nores 2004; Oyuela-

Caycedo 1996; Suguio 1993; Weiland 1984).

Drier episodes after 1150 BP coincided with increased El Niño activity (Lachniet, et al. 2004). 25

Figure 2.6: Hypothetic Parita Bay´s ancient coastline during 1100 BP

Vampiros rockshelters Background considerations

Cueva de los Vampiros was first investigated during a long-term multidisciplinary field project in the mid- 80‘s: the Proyecto Santa Maria (PSM) that carried out the research in the river basin of the same name. The archaeologists designed the project to investigate the origins of Panamanian Pre-Columbian communities addressing several hypotheses. In a small hill called Cerro el Tigre, four rockshelters were found in 1982. The same year two test pits were dug by Cooke, Weiland and Ranere (Cooke and Ranere 1984; Weiland 1984) dug two test pits in one of these shelters, Vampiros-1: one test pit-1(1 X 2m) under the overhang and the other tespit-2 (1 X 1m) was placed on the talus slope. Their initial testing reported

two phases: the first represented by diagnostic bifacial thinning flakes associated with a date of 8560  160 (Beta-5101) and arrowroot phytoliths dated to around 8560 ± 160 BP. The second phase is represented by fish bones associated with a ceramic occupation dated of

1850 100BP.

Afterwards, when the upper ceramic layers were to be disturbed to construct large shrimp tanks and roads, Richard Cooke suggested a rescue archaeological project. This project, directed by Pearson from 2002 to 2006, reopened the old test pits from 1982

(Pearson and Cooke 2002; Pearson 2002; Pearson, et al. 2003). Subsequently, Pearson and colleagues carried out extensive excavations in Vampiros-1 and Vampiros-2 and reported two fluted point fragments and a spurred end-scraper dated to 11550 ± 140 BP. This research also redefined the stratigraphic sequence (see stratigraphy below), showing that the small sample of shell and fish bone had been wrongly associated with the 8500± 160 BP date during 1982 excavations. The dated material had entered these early deposits through animal burrows from more recent ceramic deposits.

In August of 2005, I excavated two 0.5 m2 columns from the south wall of

Vampiros-1 (Figures 2.7, 2.8) and one 0.5 m2 column from the north wall of Vampiros-2

(Figures 2.9, 2.10) with the help of Lourdes Castillero, Ubencio Vargas and Lisbeth Valencia.

My purpose was to obtain samples for analysis to determine whether the two Vampiros shelters functioned as specialized fishing stations at which marine fish acquired in nearby inshore and river-mouth habitats were salted, smoked and/or sun-dried for subsequent transport to coeval communities located further inland. These column samples provided

diverse number of vertebrates, mollusks, ceramics and stone-tool artifacts which date between ca 2200 and 1150 BP.

Figure 2.7: Location of Vampiros-1. Top: Cross section of Vampiros-1 (Pearson 2002). Left: View of the east wall with the 2005 columns. Right: General view (Pearson 2002).

Figure 2.8: Map of Vampiros-1 showing excavations units 1982-2006 (TP) and column samples 2005 #1 (right) and # 2 (left).

Figure 2.9: Location of Vampiros-2.Top: Cross section of Vampiros-2 (Pearson). Left: View of the north wall with the 2005 column. Right: General view (Pearson).

Figure 2.10: Map of Vampiros-2 showing excavations units 2005-2006 (TP) and the 2005 column sample# 3. 32

Stratigraphy and Deposits

The archaeological work conducted during ―Proyecto Santa Maria‖ in 1982 (Cooke and

Ranere 1984) and that of Pearson and colleagues from 2002 to 2006 (Carvajal Contreras, et al. 2008; Pearson and Cooke 2002; Pearson 2002; Pearson, et al. 2003) identified 5 macrounits summarized as follows:

 Macrounit 5 (D5) is the basal deposit of Vampiros -1 (Figure 2.11). There is

no evidence of Cultural materials (Carvajal Contreras et al 2008; Pearson

2002).

 Macrounit 4 (D4) is a thin anthropogenic soil initially occupied during

Paleoindian times at Vampiros -1(bulk sediment date: 11550 ± 140 BP). Three projectile points, scrapers and bifacial flakes associated with Clovis-like and Fish tail traditions were found. Neither shell nor vertebrate remains were present in these deposits (Figure 2.11).

 Macrounit 3 (D3) is a band of brown soils with no cultural materials. This layer is the evidence for the abandonment of this shelter after about 7700 BP.

Pearson (2002: 69) proposes that this layer is a result of either the overflow of the Santa Maria at the base of the site or that the inselberg was once surrounded by the sea. Barber (1981:15) mentions that the main channel of the Santa Maria

River changed in the past suggesting that the Estero Salado 6.7 km north to

Cerro Tigre was the marine outlet for this river (Figures 1.1, 2.2 and 2.11).

 Macrounit 2 (D2) corresponds to a ceramic-period occupation and consists

of an array of sediments composed of charcoal, ash, marine mollusk and

vertebrate remains, shell artifacts, stone flakes and pottery as well

anthropogenic features such as postmolds and hearths (Figure 2.12). Fourteen

14C dates were obtained (Figure 2.13).

Macrounit 1 (D1) is the uppermost unit (Figure 2.12). Sedimentation at the

top of both shelters (D1) was considerably slower than in D2. This deposit

contains shell and vertebrate remains. Several dates were obtained (Figure

2.13): 1140 ± 40 BP (Beta-217529) and 1170 ± 70 BP (Beta-217530), in

Vampiros-1, and 1190 ± 40 BP (Beta-217527) in Vampiros-2. Macrounit 1 and 2 will be discussed on detail in the following section.

Figure 2.12: Stratigraphic profile of Macrounit 1 and 2, Vampiros-2 (north wall) and Vampiros-1 (east wall) (Carvajal et al 2008:95).

Figure 2.13: Radiocarbon dates for Vampiros-1 and Vampiros-2

Macrounits 1 and 2

The macrounits were divided into subunits. These divisions were defined on the basis of changes in sediment color, sediment texture, charcoal bands and artifact assemblages (i.e. frequency of pottery, stone tools artifacts, shell species and bone density). As mentioned above, Vampiros-1 was composed of 2 depositional units and 29 subunits (Figures 2.14 and

2.15). The ceramic layers at Vampiros- 2 were distributed in 2 depositional units and 59 subunits (Figure 2.16). The following Table summarizes the information. The description includes macrounit (1, 2 etc) and subunit designations (1, 2, hoyo 6, etc), Munsell soil chart color designation, the sedimentary composition of the soil and remarks.

Macrounit 1 Sediment Comments Shelter Subunit Column Color Texture 1 10yr 4.3/2 Sandy loam Soft soil, eroded shells and unburned bones. Guano 2 5yr 4/2 Sandy loam Soft soil with whitish particles. Unburned bones 3 7.5yr 4/2 Sandy loam Soft soil, hearth on column 2(?), burned bones 4 7.5yr 4/2 Sandy loam Soft soil with reddish particles. Unburned bones Vamp-1 5 c1-c2 7.5yr 4/2 Sandy loam Soft soil and ash, unburned shells and bones 1 10yr 4/2 Sandy loam Soft soil, eroded shells and unburned bones. Guano 2 10yr 5.4/2 Sandy loam Soft soil with whitish particles. Unburned bones 3 10yr 4/2 Sandy loam Soft soil with whitish particles. Unburned bones 4 10yr 3/2 Sandy loam Soft soil with whitish particles. Unburned bones 5 10yr 7/2.3 Sandy loam Soft soil with whitish particles. Unburned bones 6 10yr 7/2 Sandy loam Soft soil and laminar ashes Vamp-2 7 c3 10yr 7/2 Sandy loam Soft soil and laminar ashes

Table 2.1: Soil characteristics of the stratigraphic subunits at Macrounit 1

Figure 2.14: Detailed stratigraphic profile of column 1, Vampiros-1.

Macrounit 2a Sediment Comments Shelter Subunit Column Color Texture 6 10yr 4.3/2 Sandy loam Laminar ashes with unburned bone and shells 7 5yr 4/2 Sandy loam Loose sediment with charcoal particles and unburned bone and shells 8 7.5yr 4/2 Silty loam Loose sediment with charcoal particles and unburned bone and shells 9 7.5yr 4/2 Silty loam Loose sediment and ashes. Pit (?) Unburned bone and shells Vamp-1 Hoyo 3 c1-c2 7.5yr 4/2 Sandy loam Posthole. Burned bones and shells 8 7.5yr 3/2 Sandy loam Ashes, unburned bone and shells 9 10yr 4.3/2 Sandy loam Compact reddish sediments. Hearth(?) Unburned shells 10 2.5yr 7/3 Sandy loam Loose sediments, burned bones 11 7.5yr 3/2.3 Sandy loam Loose sediment and unburned bone and shells 12 7.5yr 3/2 Sandy loam Loose sediment with reddish particles. Burned bone and shells 13 7.5yr 3/3 Sandy loam Loose sediment with yellowish particles. Burned bone 14 7.5yr 3/3 Sandy loam Compact ashes burned bones 15 7.5yr 3/2.3 Silty loam Compact ashes burned bones 16 7.5yr 4.3/3 Silty loam Compact ashes burned bones 17 7.5yr 4/3 Silty loam Compact ashes 18 7.5yr 4/3 Silty loam Compact ashes 19 7.5yr 4/3 Silty loam Compact ashes. Dense and burned bones 20 7.5yr 3/3 Silty loam Compact ashes burned bones 21 7.5yr 3/3 Silty loam Compact ashes burned bones 22 10yr 4/3 Sandy loam Compact ashes burned bones 23 10yr 4/3 Sandy loam Compact ashes burned bones 24 2.5y 5/2 Silty loam Compact ashes burned bones 25 2.5y 5/2 Silty loam Loose ashes, unburned bones 26 10yr 6/4 Loam Compact ashes, unburned bones 27 10yr 6/4 Silty loam Compact ashes burned bones 28 10yr 6/4 Silty loam Compact ashes burned bones 29 10yr 6/4 Silty loam Compact ashes burned bones 30 10yr 6/4 Silty loam Compact ashes burned bones 31 10yr 6/4 Silty loam Compact ashes, unburned bones 32 10yr 6/4 Silty loam Laminar ashes with unburned bone and shells 33 10yr 5/4 Sandy loam Laminar ashes with reddish particles Hoyo 13 7.5yr 3/2 Sandy loam Posthole. Few bones Hoyo 15 7.5yr 3/2 Sandy loam Posthole. Few bones Hoyo 18 10yr 4/3 Sandy loam Posthole. No bones Hoyo 23 10yr 4/3 Silty loam Posthole. Burned bones and shells Vamp-2 Hoyo 27 10yr 6/4 Silty loam Posthole. No bones Hoyo 28 c3 10yr 6/4 Silty loam Posthole. Few bones Table 2.2: Soil characteristics of the stratigraphic subunits at Macrounit 2a

Figure 2.15: Detailed stratigraphic profile of column #2, Vampiros-1

Macrounit 2b Sediment Comments Shelter Subunit Column Color Texture 11 10yr 4.3/2 Silty loam Greasy sediments unburned bones 13 10yr 5/4 Silty loam Compact ashes unburned bones 14 7.5yr 4/2 Silty loam Compact ashes, charcoal particles and unburned bones 15 7.5yr 4/2 Silty loam Loose sediment with greasy sediments. Unburned bones 16 7.5yr 4/2 Silty loam Ashes with orange band. Unburned bones 17 10yr 6/3 Silty loam Compact ashes burned bones 18 10yr 6/2 Silty loam Compact ashes, greasy particles and unburned bones 19 10yr 4/2 Silty loam Compact ashes, greasy particles and unburned bones 20 10yr 4.3/2 Silty loam Laminar ashes and loose sediment. Unburned bones 21 5yr 4/2 Silty loam Compact ashes burned bones 22 7.5yr 4/2 Silty loam Compact ashes, greasy particles and unburned bones 23 7.5yr 4/2 Silty loam Compact and laminar ashes. Burned bones 24 2.5yr 8/3 Silty loam Compact ashes burned bones 25 2.5yr 8/3 Silty loam Sediment with charcoal, burned bones 26 5yr 4/2 Silty loam Compact sediment burned bones 27 10yr 6/2 Clay loam Loose sediment and unburned bones 28 7.5yr 4/1 Clay loam Compact sediment unburned bones 29 10yr 4/3 Clay loam Loose sediment and unburned bones Hoyo 5 7.5yr 3/2 Clay loam Posthole, unburned bones Hoyo 6 7.5yr 3/2 Silty loam Posthole, burned bones Vamp-1 Hoyo 10 c1-c2 7.5yr 3/2 Silty loam Posthole, burned bones Hoyo 12 10yr 5/4 Silty loam Posthole, unburned bones 34 10yr 6/2 Silty loam Compact ashes unburned bones 35 10yr 6/4 Silty loam Compact ashes burned bones 36 7.5yr 5.4/3 Clay loam Compact ashes and charcoal. Burned bones 37 7.5yr 4/2 Silty loam Compact ashes and charcoal. Burned bones 38 7.5yr 3/2 Silty loam Compact ashes and charcoal. Burned bones 39 7.5yr 5.4/3 Silty loam Greasy sediments and few bones 40 10yr 5/3 Silty loam Compact ashes . Burned bones 41 10yr 4/3 Silty loam Greasy sediments and burned bones 42 10yr 4/3 Silty loam Ashy sediments and burned bones 43 10yr 4/3 Silty loam Ashy sediments. No bones 44 7.5yr 4/2 Silty loam Compact sediment and burned bones 45 7.5yr 3/2 Silty loam Compact sediment and unburned bones 46 7.5yr 3/2 Silty loam Compact sediment and unburned bones 47 7.5yr 3/2 Silty loam Compact ashes and unburned bones 48 7.5yr 4/3 Silty loam Compact ashes and unburned bones 49 7.5yr 4/3 Silty loam Sediment with charcoal, unburned bones 50 7.5yr 3/2 Clay loam Compact ashes unburned bones 51 7.5yr 3/1 Clay loam Compact ashes unburned bones 52 7.5yr 3/2 Clay loam Sediment with shell fragments 53 7.5yr 3/2 Clay loam Compact sediment and unburned bones 54 7.5yr 3/2.3 Clay loam Compact sediment and unburned bones 55 7.5yr 3/2.3 Clay loam Compact sediments and few bones 56 7.5yr 3/2 Clay loam Compact sediments and few bones 57 10yr 3/2 Clay loam Compact sediments and few bones Vamp-2 58 c3 7.5yr 4/2 Clay loam Compact sediments and few bones Table 2.3: Soil characteristics of the stratigraphic subunits at Macrounit 2b.

Figure 2.16: Detailed stratigraphic profile of column #3, Vampiros-2 43

Figure 2.17: Geochemical analyses of Vampiros-1 (top) and Vampiros-2 (bottom).

The rate of accumulation for the whole stratigraphic sequence of Vampiros-1 was 2 mm per year and Vampiros-2 was 4 mm pear year.

As can be observed in Figures 2.14, 2.15, 2.16, 2.17 and Tables 2.1 to 2.3 at both localities, macrounit 1 is part of the modern ground surface. Pedogenic processes have perturbed the stratigraphy (Rivera, personal communication 2008 and Botero, personal communication 2007). High levels of phosphates are found in subunits closer to the surface and could be the result of the activities of recent fauna (i.e. bats, crabs and rodents) and plants. Macrounit 1 is the uppermost layer at both shelters and contains fewer stone tools and a higher density of pottery fragments than macrounit 2a. Faunal remains are scarce, and shell remains have lost their external coloration in contrast to those mollusks found in macrounit 2. Human activity diminishes in this macrounit. There is little evidence of ash lenses or hearths, and there is no evidence of postholes. Sedimentation of this macrounit in both shelters was considerably slower than in macrounit 2. This unit is a little more than 0.5 m deep in Vampiros-2 and ca. 0.3 m deep in Vampiros-1. The latest dates from D1 are 1140

± 40 BP (Beta-217529) and 1170 ± 70 BP (Beta-217530), in Vampiros-2, and 1190 ± 40 BP

(Beta-217527) in Vampiros-1. These deposits are granular in appearance and have been disturbed by natural agents. Frog and bat bones and roots were most frequent in this unit.

Macrounit 2, as mentioned above, consists of superimposed anthropogenic features such as postholes, ash floors, charcoal bands and hearths. Low levels of phosphates indicate that Vampiros inhabitants kept both shelters free of animals (Rivera, personal communication 2008). This macrounit was subdivided in two episodes in both shelters.

Human activity started at macrounit 2b and is limited to occupation floors (Vampiros-1 subunits 28 to 29 and Vampiros-2 subunits 53 to 59), post holes, charcoal bands and ash lenses. High density and diversity of faunal remains are coupled with a moderate density of pottery remains and high density of lithic artifacts .These compact deposits have a laminar

appearance and accumulated fairly slowly between 2200 and 2050 BP (Figures 2.13, 2.14,

2.15 and 2.16). The deposition of macrounit 2a was very rapid between ca. 2050 and 1920

BP (more rapid in Vampiros-2 than in Vampiros-1 see Figure 2.16). These deposits are granular in appearance and are disturbed by prehistoric human activities. Post holes are abundant in both shelters (hoyo 3, hoyo 13, hoyo 15, hoyo 18, hoyo 23, hoyo 27 and hoyo

28), as well as ash lenses, pits and hearth and less charcoal bands. This unit contains less cultural material (i.e. pottery and lithic artifacts) than macrounit 2b.

Predictably the levels of organic matter are higher in macrounit 2 than in macrounit 1 because of the great quantity of shell and vertebrate remains. The soils of both macrounits are only slightly acidic, due to intense leaching (Carvajal Contreras, et al. 2008). On the other hand, soil texture in macrounit 1 and macrounit 2 has similar matrices (sandy loam). The sand proportion in all these depositional units is quite uniform. This suggests that the soils deposited on the surface of each unit, which normal erosive processes would have made sandy, were constantly transported downwards by human and/or animal activities (Figure

2.16). The sediments came also from the dry salt flats and strong dry season winds. The greater accumulation reflects the more extensive albino formation after ca. 1900 BP.

The radiocarbon chronology based on charcoal is internally consistent, with the exception of Beta-217522. This minor reversal suggests a localized human disturbance in subunit 22, produced perhaps by a posthole (subunit hoyo 5). As observed on profiles, the strata are distributed in almost horizontal layers without severe mixing, except for local human disturbances in macrounit 2a (i.e. postholes) and animal activity on macrounit 1 such as small animal burrows from wasps, crabs, iguanas and rodents (Carvajal Contreras, et al.

2008). Temporally diagnostic artifacts such as pottery (see artifacts chapter) found in 46

macrounit 2, indicate occupations coeval with already described sites in Central Pacific

Panama with La Mula and early Aristides pottery styles dated from 200 BC to 250 AD.

Lithics from the first millennium BC were found also: three La Mula points found in macrounit 1 and macrounit 2a respectively, a blade in macrounit 2b, and celts and flakes from polished celts in macrounit 2b (Cooke and Ranere 1992b; Cooke, et al. 2000; Hansell

1988; Isaza 1993).

Field methods

Although Pearson and colleagues recorded carefully the stratigraphy of the ceramic layers when Vampiros shelters were re-excavated, they did not sample fish remains. They recovered vertebrate (no-fish) remains, and artifacts. My challenge was therefore to obtain a sample that would help me to understand temporal changes and at the same time information of discrete features related to specific fishing activities. I decided to use the exposed walls of each shelter as a guide. Starting at the top of the profiles of each shelter, I cut the sections back only half a meter and excavated downward by cultural levels corresponding to macrounit 1 and macrounit 2. I scraped the surface and plotted artifacts in situ. In Vampiros-1 both columns were 2 meters in depth, while the Vampiros-2 column reached 4.5 meters in depth. These three columns were dug to sterile soil (macrounit 3).

All the sediments were screened using 1/8‖ and 1/16‖mesh size. The sediments below

1/16‖ mesh were measured (weight and volume) and bagged in the field for transport to the

Smithsonian Tropical Research Institute´s laboratory of Archaeology at Naos. All the materials were saved. In June of 2006 I started to work with the sediment samples and the artefacts with the assistance of Alexandra Lara K., Lisbeth Valencia S. and Jacqueline 47

Sánchez. The sediments from the three column samples excavated in 2005 were water- screened in the laboratory. These samples were sieved using a fine mesh of 0.25, 0.125 and

0.0625 mm.

Each cultural level was labeled by a code. The ceramic occupational zone at Vampiros-1 was composed of 2 depositional units and 29 stratigraphic subunits. The ceramic layers at

Vampiros-2 comprised 2 depositional units and 59 stratigraphic subunits.

Chapter 3

Cueva de los Vampiros and Its Cultural -Geographical Setting

Introduction

The Isthmus of Panama connects North and South America as a passageway or landbridge.

Panama has been subject to several archaeological studies since the 19th century. Over the decades, the nature of these studies has been transformed, reflecting changes in the archaeological paradigms from historical particularist to evolutionary and ecological approaches (Cooke 2005; Cooke and Sanchez Herrera 2004; Fitzgerald 1994; Politis 2003).

Panama occupies an important place in New World archaeology. Research has yielded evidence critical to our understanding of the initial peopling and colonization of the

Americas ( Cooke 2005; Cooke and Ranere 1992c; Pearson and Cooke 2002; Pearson 2002;

Ranere and Cooke 1991, 1995, 2003), the beginnings of plant cultivation, food production, sedentism and of early pottery production (Cooke 2005; Cooke and Ranere 1989; Cooke and

Ranere 1992c; Cooke 1995, 1998b; Cooke and Ranere 1994; Dickau, et al. 2007; Griggs

2005; Linares and Ranere 1980; Perry, et al. 2007; Piperno and Holst 1998; Piperno 2006;

Piperno in press; Piperno, et al. 1992; Piperno and Jones 2003; Piperno 2000; Willey, et al.

1954). It has also been a subject of intense research related to paleoenvironmental reconstruction and human impacts on the landscape (Barber 1981; Bush and Colinvaux

1994; Clary, et al. 1984; Cooke, et al. 2007; Cooke, Sanchez, et al. 2003; Dere 1981; Piperno in press). As well, several studies have been focused on the emergence and development of

social complexity (Cooke and Ranere 1992b; Cooke 1979, 2004a, b; Hansell 1988; Isaza-

Aizpurua 2007; Lothrop 1937, 1942a; Mayo 2004; Mendizábal Archibold 2004).

Situated as it is between North and South America, archaeological studies have also grappled with defining cultural-geographic boundaries of Panama in relation to neighbouring regions. The region has been ascribed to categories such as ―Lower Central America‖, ―the

Intermediate Area‖, ―Northwest South American Littoral Tradition‖ and the ―Isthmo-

Colombian Area‖, using cultural geography (Sheets 1992; Steward 1963), linguistics

(Constela Umaña 1995; Fonseca and Cooke 1994), genetics (Arias, et al. 1992; Barrantes, et al. 1990; Ruiz -Narvaez, et al. 2005), subsistence economy or material culture as the defining criteria (Bray 1984, 1992; Cooke and Sanchez 2001; Hoopes and Fonseca 2003; Willey

1971). The country as well has been subdivided into a tripartite model based on material culture: Gran Chiriquí (Western), Gran Coclé (Central) and Gran Darien (Eastern) (Cooke

2005; Cooke 1976; Haller 2004; Isaza-Aizpurua 1993; Isaza-Aizpurua 2007; Palumbo 2009).

Cueva de los Vampiros is situated in Central Pacific Panama and part of Gran Coclé.

Figure 3.1: Map of Panama, showing archaeological sites that are discussed in this text.

Central Pacific Panama

Central Pacific Panama has received more attention from archaeologists than any of the other areas. It has a copious zooarchaeological record because the animal remains deposited in several dumps, habitation areas, burials, workshops and caves are well preserved. This area, also known as Gran Coclé, has a pre-Columbian history extending from Paleo-Indian times to the Spanish conquest ( see Table 3.1)(Bird and Cooke 1974;

Carvajal Contreras, et al. 2005; Cooke 2005; Cooke, et al. 2008; Cooke and Ranere 1999;

Cooke and Sanchez Herrera 2004; Cooke 1995; Díaz 1999; Dickau 2005; Dickau, et al. 2007;

Fitzgerald 1994; Haller 2004; Hansell 1979, 1988, 1992; Hansell 1997; Mayo and Cooke

2005; Mayo 2004; Pearson 2005; Pearson and Cooke 2002; Pearson 2002; Pearson, et al.

2003; Perry, et al. 2007; Ranere 1992; Ranere and Cooke 1991, 1995; Ranere and Cooke

1996; Ranere and Hansell 1978; Sanchez Herrera 1995; Sanchez Herrera and Cooke 2000;

Valerio 1985, 1987).

Over the last thirty years some of the multidisciplinary research has focused on the utilization of animal resources, vertebrates and invertebrates, for dietary purposes and for the creation of tools and ornaments. Although fish remains are abundant in archaeological sites as early as the Preceramic at Cerro Mangote (ca. 7000 BP), it was not until the

―Proyecto Santa Maria‖ research project (PSM) was initiated that zooarchaeological research focused on fish exploitation and fishing in prehistoric societies (Cooke and Jiménez 2004;

Cooke and Ranere 1984; Cooke and Ranere 1989; Cooke and Ranere 1992c; Cooke 1992a,

1998b, 2004a, b; Hansell 1979). Earlier ―zooarchaeological‖ studies have been centered on shell or bone objects from mortuary contexts at Sitio Conte (Lothrop 1937, 1942a). Later

archaeologists who studied shell bearing middens such as Monagrillo and the coastal site

Cerro Mangote were concerned mostly with marine mollusks as a food resource and environmental indicator (Willey, et al. 1954 ). They did not use fine mesh to recover fish bones, and it is not until the PSM that it became a standard practice. Monagrillo and Cerro

Mangote were re-excavated in 1975 and 1979 respectively by Ranere to collect better samples using multiple screens and column samples (Cooke and Ranere 1994; Cooke and

Sanchez Herrera 2004; McGimsey 1956; Ranere and Hansell 1978; Willey, et al. 1954).

During the last 30 years, improved methodologies and actualistic studies were used to analyze and reanalyze animal remains and their distributions in coastal and inland sites at

Monagrillo, Carabali shelter , Corona shelter , Cueva de los Ladrones, Aguadulce shelter,

Cerro Mangote, Sitio Sierra and Cerro Juan Diaz in order to understand other aspects of the pre-Columbian societies, such as subsistence strategies, seasonality and site location on the ancient coastline (Cooke and Jiménez 2004; Cooke, et al. 2008; Cooke 2001; Cooke and

Tapia-Rodríguez 1994a, b; Hansell 1979; Jiménez 1999; Jiménez and Cooke 2001; Zohar and

Cooke 1997).

Still more recently, zooarchaeological analyses in domestic contexts at Zapotal, Cerro

Juan Diaz and Vampiros correlated animal taxonomy and their abundance to niche exploitation, mobility, cooking /gathering methods and taphonomy (Belanger 1999; Carvajal

Contreras 1998; Carvajal Contreras, et al. 2008; Cooke, et al. 2007; Peres 2001). Other studies used animal remains from mortuary and domestic contexts to document craft specialization (Isaza-Aizpurua 2007; Lyall 2007; Mayo Torné 2004), cosmology and social complexity (Cooke 1989; Cooke 1998a, 2004a, b; Haller 2004).

Table 3.1: Chronological sequence of ―Gran Cocle‖.

Chronology

The lower ceramic-bearing layers of Vampiros are attributed to an early Formative period called the Middle Ceramic B which is based on artifact typology and radiocarbon dates. This period, between 400 B.C. to A.D. 250, is characterized as a time antecedent to the rise of chiefdom societies (Cooke 2005; Cooke and Ranere 1992b; Dickau 2005; Griggs 2005;

Haller 2004; Isaza-Aizpurua 2007; Sanchez Herrera 1995; Sanchez Herrera and Cooke 2000).

The period from around the mid-first millennium B.C. to A.D. 300, was one of rapid cultural change, which included sedentism, hierarchical settlements, agricultural intensification and craft specialization (Dickau 2005; Piperno 2006). Archaeological surveys suggest an increasing population that moved from the hamlets and rockshelters in the foothills to nucleated agricultural villages along the alluvial plains of the main rivers in the lowlands and the coast, at sites such as the earliest village La Mula-Sarigua, Sitio Sierra, Cerro

Juan Diaz, Bucaro, La India, Cañazas, El Indio, and El Cafetal (Cooke 2005; Cooke and

Ranere 1992b; Hansell 1988, 1992, 1997; Ichon 1980; Isaza Aizpurua 1993).

Although these are largely sedentary populations, different activities affected the duration of their occupations. These communities have also a pattern of transhumance (i.e. Asiento

Viejo and Asiento Nuevo ) or seasonal occupation (coastal sites are occupied during the dry season to collect and catch inshore fish)(Cooke and Ranere 1992b; Haller 2004; Hansell

1988, 1992; Hansell 1997; Ichon 1980; Isaza-Aizpurua 1993; Isaza-Aizpurua 2007; Weiland

1984).

Technological changes included the development of regional pottery styles in Central

Panama associated with the emergence of slip painting: e.g., La Mula and Aristides. Large- size vessels for liquid storage occur at this time (Cooke and Ranere 1992b; Isaza-Aizpurua

1993; Sanchez Herrera and Cooke 2000). Legless slab metates, manos, and basalt polished axes were also introduced-- stone tools typically associated with the cultivation and processing of maizeand forest clearance, which became the main staple food in the region.

Sweet potatoes, squash, manioc and, capsicum peppers and palm nuts were also consumed

(Dickau 2005; Norr 1995; Perry, et al. 2007; Piperno 2006; Ranere and Cooke 1996). Stone axes were employed to clear the gallery forests to accommodate the increase of agriculture 55

activities (Cooke 1977; Dickau 2005; Piperno 2006). Blades and stemmed flake points made of cryptocrystalline rock were also found at these village sites, with edge-wear patterns indicating that they were multipurpose tools used as scrapers, knives, perforators and gravers

(Cooke and Ranere 1992b; Ranere and Cooke 1996); Hansell personal communication

2008). It has been suggested that households from both the Caribbean and Pacific watersheds were involved in the exchange of these chalcedony tools (La Mula-Sarigua) for basalt axes as well as cured fish and salt (Cooke and Ranere 1992b; Griggs 2005; Haller 2004;

Hansell 1988).

The particular animal resources that contributed to the pre-Columbian diet depended on the ecological context of each site. White- tail deer, agouti, cottontail rabbit, armadillo, iguana, marine fish, crabs and shellfish were among the most common animals consumed.

Marine shoaling and freshwater fish were captured using perishable fishing-gear ( Cooke

2005; Cooke and Jiménez 2008; Cooke, et al. 2007, 2008; Cooke 2001; Cooke and Tapia-

Rodríguez 1994b).

These communities established formal cemeteries on the tops of small hills. These burial grounds have been identified at Cerro Juan Diaz, Sitio Sierra, La Mula-Sarigua and

Tonosi Valley sites. Analyses of their mortuary offerings suggest that these groups were egalitarian (Cooke and Ranere 1992b; Díaz 1999; Hansell 1988; Ichon 1980). Briggs (1989) suggests that status was based on age, gender and activity. Most of the offerings consist of carved stone, bone and shell jewelry and eventually, gold work; all are the result of craft specialization (Bray 1992; Cooke 1977; Cooke, et al. 1998; Cooke 2004b; Cooke and Sanchez

Herrera 1997; Ichon 1980; Sanchez Herrera and Cooke 2000). The early metalwork of Gran

Coclé was found at Cerro Juan Diaz dated to around 150 B.C(Cooke 2005; Cooke, Isaza, et al. 2003; Cooke, et al. 2000; Isaza-Aizpurua 2000).

Fishing in Central Panama

Pre-Columbian fishing in the ―Gran Coclé‖ is a local activity, characterized by an intensive exploitation of marine and freshwater fish along intertidal mudflats and the river mouths.

The patterns of fish used by Pre-Columbian inhabitants show that their diversity and abundance in Central Pacific fish assemblages can be linked to the topographic position of each site and changes in the coastline. Several articles have been published about fishing practices and fish species in Central Panama (Carvajal Contreras, et al. 2008; Cooke and

Jiménez 2008a, b; Cooke and Jiménez in press; Cooke, et al. 2008; Cooke 1992a, 1996;

Cooke and Sanchez 2001; Jiménez and Cooke 2001; Zohar and Cooke 1997). Here, I summarize very briefly the Pre-Columbian fishing practices from the Preceramic period until

1100BP mentioning date of occupation, surrounding paleo-environment, frequent marine species, the site‘s distance from the coast and objects associated with fishing practices

(Tables 3.2 and 3.3).

As I mentioned above Pre-Columbian fishing practices are affected by the access to coastal environments. The first change in drainage patterns as a consequence of eustasy and the stabilization of the worldwide seal levels around 7000 BP is related with the apparent intensification of aquatic resource use after this time and later on (Barber 1981; Clary, et al.

1984; Oyuela-Caycedo 1996; Suguio 1993; Van der Hammen 1974).

As a consequence of this worldwide event, a sequence of sites in Gran Coclé such as

Cerro Mangote 1.5 km from the coast (14C to cal. 5930–5450 B.C), Monagrillo (cal 2880

[2610] 2460 B.C.) and Zapotal(cal. 1660–1220 B.C. (Beta-20850, Cooke 1995: 173) both on the coastbear evidence of the exploitation of sea foods. These coastal native communities exploited species that frequent sandy beaches, estuaries and shallow lagoons, e.g., Pacific moonfish (Selene peruviana), thread-herrings (Opisthonema spp.), marine catfish

(e.g., Ariopsis seemanni and Cathorops furthii), Pacific bumpers (Chloroscombrus orqueta), puffer fish (Sphoeropides annulatus and Guentheridia formosa) and threadfins (Polydactylus spp.)(Cooke and Ranere 1992a; Cooke 1992a, 1995; Jiménez 2003; Jiménez and Cooke 2001; Peres 2001).

In contemporaneous rock shelters further inland such as the Aguadulce Shelter

(today located 18 km from the coast), deposits (cal. 10,869–10,408 B.C. and cal. 10879–9887

B.C. (R-24531/2 and R-24531/1; Cooke 2005: 136; Cooke and Ranere 1992c)) indicate that communities exploited species similar to the sites above: Pacific moonfish (Selene peruviana), thread-herrings (Opisthonema spp.), marine catfish (e.g., Ariopsis seemanni and Cathorops furthii),

Pacific bumpers (Chloroscombrus orqueta) and threadfins (Polydactylus spp.). Another inland site,

Cueva de los Ladrones , now 25 km from the coast, Pacific bumpers (Chloroscombrus orqueta),

Pacific moonfish (Selene peruviana), thread-herrings (Opisthonema spp.), point-nosed croaker

(Ophisocion typicus) and Pacific ilisha (Ilisha fuerthii) were recovered from the Early Ceramic levels (2550-550 BC), and finally Corona Shelter (11750–8090 B.C. (Beta-19105; Cooke

2005: 136).

, today located 55 kilometres from the Gulf of Montijo, has evidence of marine fish such as point- nosed crocker (Ophioscion typicus), catfish (Ariidae) and shells, perhaps obtained through exchange with coastal communities (Cooke and Jiménez 2004; Cooke 2001).

In la Mula- Sarigua (1050BC-AD750) a village located on the coastline, people exploited puffer fish (Tetraodontidae), marine catfish (Ariidae), jacks (Carangidae) and sharks. All of these species are commonly found near mangroves and/or at river mouths

(Cooke and Jiménez 2004; Hansell 1988).

Sitio Sierra, a later site, is a large 45ha nucleated village located on the Santa Maria

River Valley. Cooke and colleagues (Cooke, et al. 2008; Cooke and Ranere 1999; Cooke

2001) studied fish remains found there in a large refuse lens deposited ca. AD 150-450. They determined that the most frequent marine species in this sample are thread-herrings

(Opisthonema spp.), Pacific moonfish (Selene peruviana) and brassy grunt (Orthopristis chalceus).

The prevalence of thread-herrings among Sitio Sierra´s archaeofaunas indicates the growing importance of gillnets. Similar species were exploited at AG-125 dated ca 1000 BP (825 ±

60 BP (cal AD 1040-1285) (GX-25700-LS), where excavations revealed structures and dumps, mostly of Crassostrea shells.

Cerro Juan Diaz (AD250-1400), a village 4km from the coast in la Villa River´s

Valley, was used for domestic and ritual activities. Analyses of five features determined that the most frequent species exploited were marine such as Pacific bumper (Chloroscombus orqueta),(Pomadasys panamensis), Pacific moonfish (Selene peruviana), marine catfish (Ariopsis seemani and Cathorops furthii), whitefin weakfish (Cynoscion albus) and tallfin croaker

(Micropogonias cf altippinis), puffer fish (Guentheridia formosa), point- nosed crocker (Ophioscion typicus), brassy grunt(Orthopristis chalceus) but freshwater species were exploited as well

(Sternopygus, Ctenolucious) (Jiménez 1999; Jiménez and Cooke 2001).

Distance from the sea or Site river (km) Approximate dates Some Aquatic taxon Other fauna Habitat Corona 55 (Montijo Ophioscion typicus, Ariidae, shelter gulf) 10000 BP Marine shells Dasypus novemcinctus Terrestial Arius cookei, Arius seemani, Cerro Arius kessleri, Dormitator Odocoileus virginianus, Mangote 1.5 to 5.5 7000 BP to 5000 BP latifrontis, Selenaspis dowii Canis familiaris Mangrove/ Estuary

Nowadays 18 but at time site was occupied, active coastlines was several kilometres Freshwater turtles Kinosternon and Sygmodon sp, Dasypus closer and close Trachemys. Loraciidae, Ariidae. novemcinctus, Bufo sp, Aguadulce saltwater Rhamdia, Pimelodella, Hoplias Iguana Iguana,Procyon Freshwater and brackish Shelter channels 7000 to 3000 BP microlepis, Loricariids lotor, snake streams. In-shore waters Ophioscion typicus, Opisthonema Ladrones libertate, Selene peruviana, Odocoileus, Dasypus cave 25 5000 to 3000 BP Ariidae, Crabs and shells. novemcinctus Terrestial Chloroscombus orqueta, Pomadasys panamensis, Selene peruviana, Ariopsis seemani, Cathorops furthii,Cynoscion albus, Micropogonias cf altippinis, Tayasu spp, Odocoileus Opisthonema libertate, Ophioscion virginianus, Iguana Zapotal on coastline 4000 BP typicus iguana, Bufo sp Active river delta Opisthonema libertate, Selene peruviana, Chlroscombrus orqueta, Odocoyleus virigininaus, Ilisha furthii, Polydactylus Dasyprocta punctata, Lagoon, mud flat/sandy opercuaris, Cathorops fuerthii, Sivilagus, Dasypus and shallow water Monagrillo on coastline 2300 to 1800 BP Tivela sp and Ostrea spp. novemcinctus. species

Table 3.2 Archaeological sites in Central Panama: distance from the sea, dates, animal resources and environment.

Distance from the sea or Site river (km) Approximate dates Some Aquatic taxon Other fauna Habitat Dasypus novemcinctus, marine turtles, Mud flat/sandy and Caranx caballus, Chaetodipterus Odocoileus virgininaus, shallow water species Vampiros zonatus, Albula sp, Sphoeroides Iguana iguana, Procyon offshore but enter Cave 2? 2200 BP annulatus(poisoning?) lotor mangrove Dasypus novemcinctus, Odocoileus virgininaus, Iguana iguana Brotogeris jugularis, Colinus cristatus, Coccyzus minor, Crotophaga sulcirostris, Cassidix mexicanus, Bufo marinus, Ameiva Opisthonema spp, Selene peruviana, ameiva, Kinosternon spp., Freshwater and brackish Sitio Sierra 12 1900 to 1500 BP Orthopristis chalceus Chrysemys scripta streams. In-shore waters Chloroscombus orqueta, Pomadasys panamensis, Selene peruviana, Ariopsis seemani, Cathorops furthii,Cynoscion albus, Micropogonias cf altippinis, Lagoon, mud flat/sandy Cerro Juan Guentheridia formosa, Ophioscion Odocoileus virgininaus, and shallow water Diaz 4 ? 2300 to 1500 BP typicus Iguana iguana species. Estuary. Chloroscombus orqueta, Pomadasys panamensis, Selene peruviana, Ariopsis seemani, Cathorops furthii,Cynoscion albus, Micropogonias cf altippinis, Lagoon, mud flat and Guentheridia formosa, Ophioscion shallow water species. AG-125 on coastline 1000 BP typicus Estuary.

Continued

Site Method of fishing Fishing Artefacts Other Land-based fishing methods and throw-nets. Some were No artefacts have been Cerro taken by hand in desiccating attributed to fishing Mangote salt-flat pools. practices No artefacts have been Corona attributed to fishing shelter Gill nets practices Smoked fish, trade? No artefacts have been Ladrones attributed to fishing cave Gill nets practices Smoked fish, trade?

No artefacts have been Aguadulce Hook and line, throw-net attributed to fishing Wider catchment area or Shelter (atarraya?) or hand? practices Exchange networks? No artefacts have been Fine mesh gill-nets and water attributed to fishing Zapotal craft practices No artefacts have been attributed to fishing Monagrillo Gill nets and traps practices No artefacts have been Cueva de los attributed to fishing Vampiros Gill net and traps practices Purveyor of smoked fish?

Table 3.3 Archaeological sites in Central Panama: methods of fishing and fishing artefacts.

Site Method of fishing Fishing Artefacts Other

No artefacts have been Depopulated? At the end La Mula- ? Hook and line, traps and/ or attributed to fishing of 1st millennium B.C. due Sarigua net practices to the formation of alvina Deer metapodials (net Fine mesh gill-nets and water weights and needles) for gill Wider catchment area or Sitio Sierra craft nets Exchange networks? Stone net weights associated Cerro Juan Fine mesh gill-nets and water with Conte Pottery (700- Diaz craft 900 d.C) Continued

Ethnohistorical Evidence of Exchange and Curing Fish in Central Panama

The exchange of commodities in Central Panama such as polished stone tools, colorful marine shells, fish and marine mammal bones and teeth, between the Caribbean and Pacific, has been documented archaeologically back as far as Preceramic times at the site Cerro

Mangote (Cooke et al. 2003; Cooke, Sanchez, et al. 2003; Cooke 1988; Cooke and Sanchez

Herrera 1997; Cooke and Sanchez 2001; Griggs 2005; Griggs, et al. 2002) and ethnohistorical documents mentioned the exchange of sea salt, and cured fish (de Oviedo y Valdés 1535,

1849-1855[1944], 1950; Jopling 1993). Spanish chroniclers described the native communities during the 16th century as socially and politically stratified, and occupying nucleated villages distributed in the major river valleys. They were ruled by paramount chiefs named Natá,

París, Escoria, and Guarare. Social interactions between these polities, who spoke Chibchan languages, consisted of both extensive trade and warfare (Cooke 1993).

Oviedo describes peaceful periods between wars when inland and coastal communities exchanged products. They traded many commodities, including salted fish, either by boat or on the backs of slaves (see Figure 3.2):

Figure 3.2: Native Americans transporting goods and crossing a river in Costa Rica (Ibarra

1999: 67)

―cuando no están en la guerra todo su ejercicio es tratar y trocar cuanto tienen unos con otros. Y así, de unas partes a otras, los que viven en las costas de la mar o por los ríos, van en canoas a vender de lo que tienen complimiento y abundancia y a comprar lo que les falta. Y asimismo tratan por tierra y llevan sus cargas a cuestas de sus esclavos: unos llevan sal, otros maíz, otros mantas, otros hamacas, otros algodón hilado o por hilar, otros pescados salados‖ (Cooke and Sanchez 2001; de Oviedo y Valdés 1849-1855[1944]: 40; Ibarra 1999).

There are few references about fish, fishing practices or curing methods. Among them, Espinosa describes people called choriga (subordinate?) from the coast of Parita Bay who exchanged fish and crabs for maize at Natá.

―… En este tiempo iban y venian muchos indios chorigaras, con cangrejo y pescado a rescatar maíz al real de manera que andaban por las calles del real vendiendo su mercancía…‖ (Carvajal Contreras, et al. 2008; Jopling 1993:49).

Carl Sauer (1966: 274), citing Spanish chroniclers, says that Cueva people preserved fish, and game by using drying racks (barbacoas) and salting, drying and smoking. The salt was provided by Chiru, Nata and Paris chiefdoms.

Inland transport of marine fish was also mentioned for Comogre´s territory in

Western Panama. According to Andagoya, people living on the Pacific slopes obtained fish from the Caribbean coast:

―…está un cacique que se dice Comogre y otro que se dice Pocorosa, están tan cerca de la mar el uno como el otro; tienen mucho guerra unos con los otros, en toda la tierra tiene cada uno dellos un pueblo y dos a la costa de este mar, de donde se mantienen de pescado la tierra adentro…‖ (Cooke and Sanchez Herrera 2004 ; Cooke and Sanchez 2001; Jopling 1993: 24).

Oviedo (1535: capítulo XXXVII) describes how native people in Darien (Eastern

Panama) cured fish without salt and how they used the dry conditions, digging a shallow hollow in the ground, covering it with earth and burying the fish for five or six days:

―Este depósito o nueva leción, me paresce que es una cosa no oída ni vista antes, ni escripta de otra provincia alguna de la forma que en la costa de Sanct Miguel, en la Nueva Castilla, los indios adoban el pescado e lo hacen cecial, sin le echar sal, Y es desta manera: abren el pescado, e cavan en tierra hasta un palmo en hondo, e cúbrenlo allí de tierra, e está así enterrado cinco o seis días, e a cabo dellos sácanlo curado, e sale mejor que el muy buen pescado cecial de Galicia o Irlanda, e tan enjuto; e se tiene después, así, todo el tiempo que quieren, Esto se hace donde he dicho, en la cual tierra nunca llueve; e a donde adoban e curan el pescado como está dicho, es apartado de la costa de la mar cincuenta pasos más o menos…‖

Andagoya (Jopling 1993: 32) did not mention the use of salt to preserve fish either.

However at the Santa Maria River, he mentions that in chief Escoria´s territory there were excellent salt-pans produced during the dry season:

―…que tenia sus poblaciones en un rio grande ocho leguas de Meta. Aquí había muy grandes y hermosas salinas, que se cuaja de verano."

Balboa suggests that inland chiefdoms such as Comogre and Pocorosa controlled fishing resources on the Caribbean Sea that supplied their own village ―cotos‖ (Jopling 1993:

24):

―…entrando la tierra adentro hasta doce leguas, está un cacique que se dice Comogre y, otro que se dice Pocorosa, están tan cerca de la mar el uno como el otro; tienen mucha guerra con los otros, en toda la tierra tiene cada uno dellos un pueblo y dos a la costa de este mar, de donde se mantienen de pescado la tierra dentro.‖

This control over fish resources may also have occurred on the Pacific Coast (Cooke and Sanchez 2001; Helms 1979).

There is no specific information of markets or trade specialists, however as I mentioned above, Espinosa describes that the main bohío (chief´s house?) at Natá in Central

Pacific Panama functioned as a market for trade and exchange of maize, fish and crabs

(Espinosa 1994a:49). On the other hand, ethnographic descriptions mention an indigeneous ritual exchange ceremony called balseria which is practiced by the modern Native

Panamanian Ngöbé in Chiriqui. During this ceremony, two communities are involved on ritualized warfare, feasting, drinking of chicha and the exchange of crafts and goods (Young

1976, 1980a, 1980b). Helms (1979:34) argues that a ceremony like balseria could have be used by Panamania chiefdoms tied into long-distance exchange networks to obtain exotic goods, gold and esoteric knowledge.

To sum up, Panama, particularly the Central region, has been subject to several studies that increase our knowledge of endogenous social processes and change in the

Americas. Zooarchaeological research has documented the subsistence strategies of past peoples. Specifically, native people on coastal sites of Gran Coclé had exploited intensively the fishing resources since Preceramic times. Central Pacific sites exploited small shoaling fish, caught in large numbers in intertidal traps or gillnets. Archaeological and ethnohistorical data sustain that highland communities obtained cured fish by exchange with coastal communities.

Chapter 4

Pottery, Shell and Lithic Artefacts at Cueva de los Vampiros (AG-145)

Introduction

The cultural assemblage at Vampiros rockshelters came from excavations conducted between 2002 to 2006 field seasons. The artefacts consisted of 1100 pottery fragments, 162 stone tools and 1 shell ornament.

Stone tools

I analyzed the few (162) stone tools using criteria such as raw material, manufacturing techniques, form, period and function (Andrefsky 1998; Ranere and Cooke 1996). As shown in Table 4.1, the lithic assemblage includes both chipped and polished stone tools.

Unmodified Core Flake Blade Shatter Eolith Point Scraper Ground and cobbles but Polished used stone tools

28 3 73 2 26 1 3 1 29

Table 4.1: Types of Lithic Materials at Vampiros-1 and Vampiros-2

This assemblage includes finished and reworked tools such as blades, points, celts, cobbles with use-wear, pebble polishers, cores and 73 flakes. Preliminary observations of this small assemblage revealed a diversity of lithic material. The analysis identified 8 types of lithic materials (Table 4.2). Vampiros inhabitants chose coarse and strong raw material

such as volcanic rocks and basalt to make ground or polished stone tools. In contrast, chipped stone tools were elaborated from cryptocrystalline jaspers.

and Core Total Point Flake Blade Eolith Shatter Scraper Material Ground Polished Unmodifi ed cobble

Andesit e 2 2

Basalt 5 7 5 17

Chert 1 1 Jasper- yellow 2 2 4

Jasper 5 10 15 Chert- red 1 2 3 Chert brown 1 1 Chert pink 1 1 Sedime ntary rock 1 1 Volcanic rock 2 1 1 9 13 Undeter mined 26 1 58 1 5 13 104

Total 28 3 73 2 1 24 3 1 27 162

Table 4.2: Types of materials at Vampiros rockshelters.

Some of the materials have patina on their surfaces which impeded identification of their source.

Chipped stone tools

The unifacial chipped stone tools comprise 3 cores, 26 shatter, 73 flakes, 2 blades, 3 points, and 1 scraper (see Table 4.3).

As Table 4.3 indicates 75% correspond to complete flakes. Cortex is rarely present on lithic fragments and most of the samples consist of tertiary flakes (57%).

Completed Fragmented Primary flakes Secondary flakes Tertiary flakes flakes flakes 55 18 6 25 42

Table 4.3: Type of flakes at Vampiros -1 and Vampiros-2

Certain features on the flakes described in Table 4.4 such as the plane striking platform, and diffuse and prominent bulbs, ripple marks and feather termination suggest that

Vampiros inhabitants were using soft hammers and hard hammers respectively to detach the flakes from the core (Winchkler 2006).

Striking platform Impact Termination

 Plane 21  Salient bulb 21  Feather 34  Cortex 11 and  Hinge 31  Line 5 prominent  Outrep 8 13  Point ripple marks assé 14   Irregular Diffuse bulb 3 and not  Concave 6 prominent 27  No ripple marks observed  Salient bulb

and not prominent ripple marks 23  Diffuse bulb

and

prominent

ripple marks

Table 4.4: Type of flakes platforms at Vampiros rockshelters.

Figure 4.1: Left, tertiary flake with use-wear. Right, Flake from a celt.

Only 9 flakes present show use-wear on the edges (Figure 4.1). The amount of tertiary flakes and shatter indicates late stages of reduction. For example, celt-flake specimens were broken off from a finished celt when it was resharpened (Figure 4.1).

Two unifacial Mula points were made of yellow jasper and the other was made from an unknown volcanic source. Both present evidence of use (Figure 4.2).

Figure 4.2: Left, La Mula point made of yellow jasper, Right, La Mula point made of unknown material.

One tertiary flake and two blades were made of red chert. Both the flake and blades have evidence of one edge being used. One tertiary flake was made of brown chert and presents potlid scars (Figure 4.3).

Figure 4.3: Left, Blade fragment. Center, blade with use- wear. Right, tertiary flake with potlids scars. 74

Finally, a broken unifacial scraper made of pink chert has a flat side, lateral retouch and has evidence of being used (Figure 4.4).

Figure 4.4: Unifacial scraper

Ground and polished Stone tools

These tools were made of basalt, andesite or volcanic cobbles, intentionally manufactured or shaped through use. Pitted surfaces and striation marks were observed in celts, cobbles with use wear, cobble pestles, pounding stones and pebbles with a flat surface. Future starch and phytolith studies would determine which plants were being processed on these tools (Table

4.5).

Polished Cobbles Pebble Hammerstones Cobble Pounding/mashing Pebble celt with polisher pestles stones with flat use- a surface wear

3 7 2 4 4 5 4

Table 4.5: Types of Ground and polished artefacts.

The complete and fragmented celts show battered marks on their surface. Either the celts were reused as hammers or were being resharpened to work wood (Figure 4.5).

Figure 4.5: Celt found at Vampiros-1

These celts are made of basalt, which is available in the vicinity of Vampiros shelters.

Similar artefacts were found at the contemporaneous site Sitio Sierra. Cooke (1977) suggests that celt and axe raw material was obtained from basalt cobbles at the Santa Maria River.

However, some reworked material made of basalt may imply that the raw material was obtained somewhere else. For example, during the ―Proyecto Santa María‖ basalt axe-

preparation workshops were found in the foothill and cordillera zones (Cooke and Ranere

1992b; Griggs 2005).

Finally, 28 pebbles without use-marks were found at Vampiros shelters. They were brought either from the beach or the Santa Maria River and they were probably used as sling stones to hunt birds or iguanas (Figure 4.6).

Figure 4.6: Unmodified pebbles

Possible Function of Stone Tools

The chipped stone tools, especially La Mula points, suggest that Vampiros inhabitants used them as knives for a variety of tasks such as cutting and scraping fish, perhaps. Polished celts may have been used to build drying racks to preserve fish (Cooke 1977; Cooke and

Ranere 1992b).

At the time of the Contact, Oviedo (1950: 118) mentions that Cueva Indians in

Eastern Panama butchered animals using stone tools and constructed structures called

barbacoas to cure fish and deer. The following quote also emphasizes that any meat has to be cure the same day that the animal was killed:

“...y después de muertos, como no tienen cuchillos para los desollar, cuartéanlos y hácenlos partes con

piedras y pedernales, y asánlos sobre unos palos que ponen, a manera de parrillas o trébedes, en un

hueco, que ellos llaman barbacoas, y la lumbre debajo, y de aquesta misma manera asan el pescado;

porque como la tierra está en clima que es naturalmente calurosa, aunque es templada por la

Providencia divina, presto se daña el pescado o la carne que no se asa el día que

muere.”

Both polished stone celts and unifacial points are also chronological markers of these deposits for the period between 2300 to 1750 B.P. The raw material could be trade with communities on the foothils (Cooke and Ranere 1984). Similar examples are observed at La

Mula-Sarigua (Cooke and Ranere 1992b; Hansell 1988) and Sitio Sierra (Cooke 1977; Isaza-

Aizpurua 1993).

Shell ornaments

One shell ornament was recovered in 2005 (Figure 4.7) near the bottom layers of Vampiros-

1. It is a spiral bead, 3.5 cm in diameter, made of Conus patricius with a biconic perforation near the edge. Similar beads are described for Cerro Juan Diaz (Mayo and Cooke 2005; Mayo

2004Lámina 25:326).

Figure 4.7: Shell ornament found at Vampiros-1

Pottery

The 1100 pottery sherds at Vampiros rockshelters were not formally analyzed. However, some preliminary data are provided such as chronological and the stratigraphic position of these materials.

The sherds are typical of the Gran Coclé pottery tradition during the Middle

Ceramic period associated to La Mula Complex which includes Aristide and la La Mula

styles (Escotá, painted group and plastic decorated group types). Based on radiocarbon

dates the activities occurred between 2300 to 1750 B.P (Table 4.6 and Figures 4.8 and 4.9)

(Carvajal Contreras, et al. 2008). The pottery found at Vampiros shelters probably served

as containers and/or serving vessels for meals such as collared jars and open bowls and

related activities during fish processing. In contrast, the vessels identified on habitational

and mortuary contexts at Sitio Sierra and also associated to La Mula Complex comprise several types: Escotá, spherical collared jars and incurved bowls; Girón, bowls or sub- globular collared jars, and Cocobó, open bowls with interior decoration, La Mula trychrome large globular jars with open collars and La Mula urns (Cooke and Ranere

1984; Hansell 1988; Isaza Aizpurua 1993, 2007; Sanchez Herrera 1995; Sanchez Herrera and Cooke 2000).

Table 4.6: Radiocarbon dates from Vampiros-1 and Vampiros-2.

Figure 4.8: Comparison painted sherds (Las Huacas, La Mula-Sarigua, Sitio Sierra (Isaza Aizpurua 1993), Vampiros (Drawing by Carvajal Contreras and Sanchez Herrera).

Figure 4.9: Comparison plastic decorations Cerro Juan Diaz (Sanchez Herrera 1995), La Mula- Sarigua (Cooke and Ranere 1992b), Cueva de los Vampiros (Drawings by Carvajal Contreras)

Preliminary observations by Cooke suggest that some of the painted motifs, as for example the one shown in Figure 4.8, resemble designs on vessels found at La Mula- Sarigua (Cooke, et al. 2007; Cooke and Ranere 1992b: Figure 9), Las Huacas and Sitio Sierra (Isaza-Aizpurua

1993).

Plastic decoration shown on Figure 4.9 are are also similar to fragments reported at la

Mula- Sarigua site (Cooke and Ranere 1992b: Figure 8), Cerro Juan Diaz (Sanchez Herrera

1995) and Sitio Sierra (Isaza-Aizpurua 1993). Some Vampiros ´pottery sherds have burnished incisions on the exterior surface. Similar decoration was reported from Cerro

Largo (Biese 1967), Sitio Sierra (Isaza-Aizpurua 1993), La Mula-Sarigua (Cooke and Ranere

1992b; Hansell 1988) and Carabali rockshelter (Mayo 2006; Valerio 1987).

Pottery sherds and site formation

An exercise of refitting pottery sherds was performed to provide some insights into the formation process of the Vampiros deposits. Basically, the analysis of horizontal and vertical sherd dispersal is intended to document the processes that affected the development of the site. Vertical and horizontal provenience was plotted using SURFER software (Bollong

1994).

Contour plots of Vampiros-1 and Vampiros-2 (Figures 4.10 and 4.11) illustrate overall trends of sherd dispersion. Movements that were largely horizontal (marked with letter ―H‖) and vertical (marked with letter ―V‖) identify the separations between sherds.

Red lines indicate an example of vessel´s dispersion.

The contour and mesh illustrations (Figure 4.10) indicate two ―refitting zones‖. For

Vampiros-1 there are: one between 30 to 80 centimeters b.s. and another between 170 to

190 centimeters b.s. For Vampiros-2 there are: one between 20 to 100 centimeters b.s. and another between 280 to 450 b.s. vertical. Noticeably, no refittings were found between 100 and 280 centimeters b.s. on Vampiros-2 (Figure 4.11), and only one refitting was found between 85 to 165 centimeters b.s on Vampiros-1. Conjoinable artifacts have more vertical distance in Vampiros-2 than Vampiros-1 as a result of differences in sedimentation rate.

A possible explanation of the difference between Vampiros-1 and Vampiros-2 vessel dispersion patterns may be that it represents several discard events or cleaning activities on the living areas and high pre- or post-depositional disturbance when people were trampling, or digging to insert support posts for curing fish. It may be the result of burrowing activities by frogs, rats and wasps as well (Carvajal Contreras, et al. 2008).

Figure 4.10: Top: Contour drawing refitting at Vampiros-1. Bottom : Mesh drawing refitting zones at Vampiros-1

Figure 4.11: Contour drawing refitting at Vampiros-2 Bottom : Mesh drawing refitting zones at Vampiros-2

Summary

The material culture fount at Cueva de los Vampiros provided data to interpret the site at several levels.

Chronologically, pottery and artefacts are in concordance with the radiocarbon dates.

This means that people who used ‗La Mula‘ and Aristides style pottery and La Mula points inhabited both shelters simultaneously.

Taphonomically, the vessel dispersion pattern might represent multiple events and/or pre- or post-depositional disturbances when people were using these shelters, as well as post-occupation animal burrowing activities.

None of the stone tools were modified for use as net weights. Vampiros stone tool assemblage, as well as other lithic assemblages in Central Panama, lacks fishing devices (net sinkers, hooks), except for Cerro Juan Diaz site where net sinkers were found associated with a shell workshop dated to ca. AD 700-900 and Sitio Sierra where net weights and needles from deer metapodials were found on the house floors cal. 65 B.C. and A.D. 475 (Cooke 1992a; Cooke and Tapia-Rodríguez 1994a; Isaza Aizpurua 1993; Mayo

2004). Perhaps these tools were made from perishable materials (i.e. wood, cotton, and bottle gourds). However, blades and projectile points may have been used to butcher fish, and polished stone tools were probably used to build drying racks to cure fish. The raw material for the stone tools was probably obtained through exchange with communities located on the foothills.

Chapter 5

Mollusc Remains at Cueva de los Vampiros (AG-145)

Introduction

Excavations at the Vampiros rockshelters produced a large quantity of invertebrate as well as vertebrate specimens. The molluscs analyzed, as with the vertebrate remains, are from the three 0.5 m2 column samples and were recovered using fine-mesh screens and analyzed following methods common to zooarchaeology (Reitz and Wing 2008).

The purpose of the invertebrate analysis is to examine the use and procurement of shellfish and the formation processes at the rockshelters at the time of its occupation. The methods focus on identifying and quantifying species present and determining taphonomic processes.

All bivalves and univalves (gastropods) from the three column samples were used for taxonomic composition and taphonomic study. I analyzed the univalves and bivalve remains sieved over a 4 mm mesh using a comparative collection of modern molluscs housed at the

Smithsonian Tropical Research Institute in Panama and referring to malacological literature such as Cruz-Soto and Jiménez (1994), Keen(1971) and Skoglund (2001, 2002) as guides for identification of molluscs of the Panamic Province.

The quantification of data in the present study includes MNI (minimal number of individuals) and NISP (number of identified specimens) for every taxonomic group found in each stratigraphic unit and provides information related to relative frequency of taxa, dietary contributions and procurement strategies (Claassen 1998; Giovas 2009; Reitz and Wing

2008). NISP (Number of identified specimens) records the number of diagnostic fragments and whole bivalves and univalves (gastropod bodies). MNI (Minimal number of individuals) is estimated for bivalves by counting left and right or upper and lower valves and paired specimens and taking the highest of the three numbers. MNI for gastropods is estimated from diagnostic morphological features such as the apex, columella or opercula counts and taking the highest number.

Similar to vertebrate remains, NISP for invertebrates is affected by differential recovery, aggregation, cultural practices, differential preservation, and sample size among other problems. MNI for invertebrates as well is influenced by sample size, and difficulties in its calculation (Giovas 2009; Glassow 2000; Grayson 1984; Jerardino 1997).

Both bivalves and univalves were weighed in grams by sample and taxon including fragments that were not considered diagnostic. Taphonomic processes were recorded such as completeness, encrustation, perforation, abrasion, and acid dissolution; cultural processes were also recorded, e.g, heating (Claassen 1998; Morrison and Cochrane 2008; Rick, et al.

2006). Measurements of complete specimens were taken to explore the mode of collecting and identifying heavy human predation (Claassen 1998; Jerardino 1997; Mannino and

Thomas 2001). All information was tabulated in EXCEL and compared by macrounits by shelter.

Habitats used for shellfish collecting

Examination of the shellfish remains at Vampiros rockshelters gives insight into which species and habitats were exploited. The invertebrate remains from Vampiros-1 and

Vampiros-2 contain marine bivalves and gastropods from 36 families, 42 genera and 54 species, as well as crab and land snails which were not analyzed in this thesis due to time constraints. The shell sample from the three columns includes 12,729 (NISP), weighing a total of 65671.4 grams. As indicated in Figures 5.1, 5.2 and 5.3, the following marine shellfish species were the most frequently consumed in the Vampiros shelters: the gastropods rock snail (Thais kiosquiformis), moon snail (Polinices cf otis) and single-banded moon snail( Natica unifasciata) and the bivalves littleneck (Protothaca asperrima), pitar venus clam (Pitar paytiensis), grand ark (Grandiarca grandis), mussel (Mytella sp), lacerate tellin(Tellina laceridens), black ark (Anadara tuberculosa), Dunker´s dosinia (Dosinia dunkeri) and pacific calico scallop (Argopecten ventricosus).

Figure 5.1: Percentage of mollusks remains in macrounit 1 at Vampiros-1 and Vampiros-2 by weight

Figure 5.2: Percentage of molluscs remains in macrounit 2a at Vampiros-1 and Vampiros-2 by weight

Figure 5.3: Percentage of molluscs remains in macrounit 2b at Vampiros-1 and Vampiros-2 by weight

The species most frequently represented at both shelters corresponds to intertidal habitats. Through macro unit 1 the shelters are dominated by the species black ark (Anadara tuberculosa), Cortez oyster (Crassostrea cortiziensis), rock snail (Thais, kiosquiformis), pitar venus clam (Pitar paytiensis), and littleneck (Protothaca asperrima ) (Figure 5.4). Sand bars at low tide were also available to exploit grand ark (Grandiarca grandis), as represented at Vampiros.

These species became less common in macro units 2a and 2b, located at the bottom of the stratigraphic sequence. The habitats most frequently represented in Macrounit 2a are the intertidal mudflat, low intertidal zone, and moderately shallow waters that included the species single-banded moon snail (Natica unifasciata) and littleneck (Protothaca asperrima).

Mussels (Mytella sp.), Dunker´s dosinia (Dosinia dunkeri) and pacific calico scallop (Argopecten ventricosus) are present in Macrounit 2a but are more frequent in Macrounit 2b, suggesting a clearer water environment dominated by gravel or sand in strong currents. The gastropod single- banded moon snail (Natica unifasciata) from the low intertidal zone is present in throughout the stratigraphic sequence of both shelters (Figure 5.5). There is a noteworthy absence or low frequency of shell species from sandy beach environments such as the clams

Donax sp and Tivela sp and brackish water environments such as marsh clam (Polymesoda boliviana).

Figure 5.4 : Exploited habitats at Vampiros-1 (MNI)

Figure 5.5 : Exploited habitats at Vampiros-2 (MNI)

Species selection

I applied a diversity index and equitability to quantify the taxonomical diversity and species richness in the macrounits of both shelters (Reitz and Masucci 2004; Reitz and Wing 1999b).

Diversity and equitability allows to evaluate subsistence strategies in terms of the variety of animal used by pre-Columbian inhabitants and the ―the evenness with which these resources are used‖ (Reitz and Massuci 2004:79). Differences in these two indexes are result of natural environment, human agency, season of site occupation, and social positioning (Reitz & Wing

2001). Both indexes are calculated for MNI. Mollusk data from Vampiros shelter are presented in the Tables 5.3 to 5.6.

Diversity is calculated using Shannon‘s Index(Reitz and Masucci 2004; Reitz and Wing

2008) where :

´ ni The number of individuals in species i; the abundance of species i.

S The number of species.

N The total number of all individuals

pi The relative abundance of each species, calculated as the proportion of individuals

of a given species to the total number of individuals in the community:

Equitability is obtained using ´the Shannon‘s function as S is the natural log of the number of species for which MNI was estimated(E. J. Reitz and M. Masucci 2004: 79).

V´= H´/loge S 98

Vp1 MNI #Taxa Diversity Equitability M1 683 31 1.169 0.179 M2a 667 30 0.758 0.117 M2b 1540 44 0.127 0.017

Table 5.1: Mollusc Diversity and Equitability at Vampiros-1

In Vampiros-1 (Table 5.1), shell remains in Macrounit 1 (M1) had 31 taxa (MNI 683);

Macrounit 2a (M2a) had 30 taxa (MNI 667); and Macrounit 2b (M2b) had 44 taxa (MNI

1540).

Vp2 MNI #Taxa Diversity Equitability M1 601 20 1.272 0.199

M2a 542 24 0.883 0.140 M2b 565 31 0.204 0.032

Table 5.2: Mollusc diversity and Equitability at Vampiros-2

For Vampiros-2 (Table 5.2), shell remains in Macrounit 1 had 20 taxa (MNI 601);

Macrounit 2a had 24 taxa (MNI 542); and Macrounit 2b had 31 taxa (MNI 565).

In terms of MNI, diversity and equitability are low at the beginning of the occupation for both shelters (Macrounit 2b) and slightly increase towards the end of the occupation

(Macrounit 1). This reflects an increasing utilization of several species; consequently,

Vampiros people were focusing their shell gathering activities on selected several taxa. In other words, Pre-Columbian inhabitants utilized almost all the molluscs available.

Dominance and Contribution to Diet

Gastropods dominate the invertebrate sample of both shelters (Tables 5.3 to 5.6). At

Vampiros 1 macrounit 1, single-banded moon snail (Natica unifasciata) accounts for 5.2%

(MNI 286) , 7.2% (MNI 396) in Macrounit 2a, and 14.7% (MNI 803) in Macrounit 2b. A similar pattern is observed at Vampiros 2 where single-banded moon-snail (Natica unifasciata) is represented by an MNI of 215 individuals (8.4%) at Macrounit 1, 14.7% (MNI 377) at

Macrounit 2a, and 12% (MNI 309) at Macrounit 2b.

The second gastropod of importance is the rock snail (Thais kiosquiformis), which is represented at Vampiros 1 by an MNI of 132 (2.4%) at Macrounit 1, 1.7% (MNI 94) at

Macrounit 2a and 5.5% (MNI 299) at Macrounit 2b. Remains of rock snail (Thais kiosquiformis) at Vampiros 2 are 2.5% (MNI 63) at Macrounit 1, 1.1% (MNI 28) at Macrounit

42a and .5% (MNI 12) at Macrounit 2b (Tables 5.4 and 5.6). Most of the remains of rock snail (Thais kiosquiformis) are fragmented.

The third gastropod in importance is the moon snail (Polinices uber). It is represented at Vampiros-1 by an MNI of 1(less than 0.1%) at Macrounit 1, (0.1% (MNI 8) at Macrounit

2a and 0.3% MNI 16) at Macrounit 2b. Remains of this gastropod are distributed at

Vampiros-2 as follows: 0.3% (MNI 7) at Macrounit 1, 0.6 % (MNI 16) at Macrounit 2a and

0.2% (MNI 4) at Macrounit 2b

100

The fourth gastropod in importance is the moon snail (Polinices cf otis). In Vampiros-1 is represented in Macrounit 1 by an MNI of 6 (0.1%), an MNI of 4 (0.1%) in Macrounit 2a and 0.3% (MNI 17) at Macrounit 2b. In Vampiros-2 it is not present in Macrounit 1 but

(0.2% (MNI 6) at Macrounit 2a, and 0.1% (MNI 3) at Macrounit 2b.

For the bivalves, the littleneck (Protothaca asperrima) is the most abundant in both shelters (Table 5.3). In Vampiros -1there is 1.4% (MNI 74) at Macrounit 1, 1.2% (MNI 65) at Macrounit 2 and 0.4% (MNI 21) at Macrounit 2b. In Vampiros- 2 there is 9% (MNI 72) at Macrounit 1, 2.1% (MNI 14) at Macrounit 2a, and 0.2% (MNI 2) at Macrounit 2b (Tables

5.5).

The second bivalve of importance is the black ark (Anadara tuberculosa). In Vampiros-

1 there is 9% (MNI 50) at Macrounit 1, 0.3% (MNI 14) at Macrounit 2a and 0.1% (MNI 5) at Macrounit 2b. In Vampiros- 2 there is 2.8% (MNI 72) at Macrounit 1, 0.9% (MNI 24) at

Macrounit 2a and 0.6% ( MNI 16) at Macrounit 2b (Tables 5.3 and 5.5).

The third bivalve of importance is the grand ark (Grandiarca grandis). In Vampiros-1 there is 0.5 %( MNI 28) at Macrounit 1, 0.5% (MNI 30) at Macrounit 2a and 0.5% ( MNI

29)at Macrounit 2b. In Vampiros-2 there is 3.3% (MNI 84) at Macrounit 1 1% (MNI 26) at

Macrounit 2a and 0.4% (MNI 11) at Macrounit 2b (Tables 5.3 and 5.5).

The fourth bivalve of importance is the pitar venus clam (Pitar paytensis) . In

Vampiros-1 there is 0.9% (MNI 51) at Macrounit 1, 0.2 % (MNI 10) at Macrounit 2a and

0.7% (MNI 37) at Macrounit 2b. On the other hand Vampiros-2 there is 1.8% (MNI47) at

Macrounit 1, 0.9% (MNI 22)at Macrounit 2b and 1.6%(MNI 42)at Macrounit 2b.

101

The fifth bivalve of importance is the Dunker´s dosinia (Dosinia dunkeri). In

Vampiros-1 there is 0.1 %( MNI3) at Macrounit 1, 0.1% (MNI 4)at Macrounit 2a and 0.6%

(MNI31) at Macrounit 2b. In Vampiros-2 this bivalve is 0.2% (MNI 5) at Macrounit 1, 0.3%

(MNI 7) at Macrounit 2a and 0.1% (MNI 3) at Macrounit 2b.

The sixth bivalve of importance is the lacerate tellin (Tellina laceridens). In Vampiros-1 there is 0.1% (MNI4) at Macrounit 1, 0.1% (MNI 3 ) at Macrounit 2a and 0.4%( MNI 20) at

Macrounit 2b. The same pattern is observed at Vampiros-2. There is 0.1 %( MNI 4) at

Macrounit 1, 0.1%(MNI 3 ) at Macrounit 2a and 0.4% (MNI 20) at Macrounit 2b.

The seventh bivalve of importance is the Cortez oyster (Crassostrea cortiziensis) . In

Vampiros-1 there is 0.3% (MNI 18) at Macrounit 1, 0.1 % (MNI 3) at Macrounit 2a and

1.1% (MNI58) at Macrounit 2b. This oyster is distributed in Vampiros-2 by an MNI of 8

(0.3%) in Macrounit 1, and an MNI of 2 (0.1%) in Macrounit 2b.

Other bivalves and gastropods species are represented in these samples by less than

1% in all macrounits (see Tables 5.3 to 5.6). Mussels (Mytella sp) and pacific calico scallops

(Argopecten ventricosus) are frequent at both shelters in Macrounit 2a and 2b. In Vampiros-1 the pacific calico scallop is distributed by an MNI of 2 (less than 0.1%) in Macrounit 2a and an MNI of 86 (1.6%) in Macrounit 2b. This scallop is represented in Vampiros-2 only in

Macrounit 2b by and MNI of 136 individuals (5.3%).

Mussels are distributed at Vampiros-1 as follows: an MNI of 9 (0.2%) in Macrounit 1 and MNI of 15 (0.3%) in Macrounit 2a and an MNI of 61 (1.1%) in Macrounit 2b. A Similar

102

situation is observed at Vampiros-2 where this mussel is represented by an MNI of 31

(1.2%) in Macrounit 1, an MNI of 9 (0.4%) in Macrounit 2a and an MNI of 68 (2.7%) in

Macrounit 2b.

103

1 2a 2b 1 2a 2b Taxonomy MNI % MNI % MNI % TotalMNI NISP % NISP % NISP % TotalNISP Anadara perlabiata 1 0.0% 0.0% 0.0% 1 1 0.0% 0.0% 0.0% 1 Anadara similis 0.0% 1 0.0% 2 0.0% 3 0.0% 1 0.0% 2 0.0% 3 Anadara sp or Grandiarca sp 77 1.4% 69 1.3% 106 1.9% 252 80 0.9% 72 0.8% 113 1.3% 265 Anadara tuberculosa 50 0.9% 14 0.3% 5 0.1% 69 101 1.1% 24 0.3% 9 0.1% 134 Andara esmeralda 1 0.0% 0.0% 0.0% 1 1 0.0% 0.0% 0.0% 1 Grandiarca grandis 28 0.5% 30 0.5% 29 0.5% 87 59 0.7% 55 0.6% 57 0.6% 171 Argopecten sp 0.0% 3 0.1% 340 6.2% 343 0.0% 3 0.0% 471 5.2% 474 Argopecten ventricosa 0.0% 2 0.0% 86 1.6% 88 1 0.0% 3 0.0% 142 1.6% 146 Carditamera affinis 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 0.0% 1 Carditamera radiata 1 0.0% 0.0% 1 0.0% 2 1 0.0% 0.0% 1 0.0% 2 cf Chione 0.0% 0 0.0% 0 Chione obliterata 0.0% 0.0% 2 0.0% 2 0.0% 0.0% 4 0.0% 4 Chione sp 0.0% 0.0% 0 0.0% 0.0% 0 Chione subrugosa 6 0.1% 2 0.0% 2 0.0% 10 10 0.1% 4 0.0% 5 0.1% 19 Corbula sp 0.0% 1 0.0% 1 0.0% 2 0.0% 1 0.0% 3 0.0% 4 Crassostrea corteziensis 18 0.3% 3 0.1% 58 1.1% 79 42 0.5% 7 0.1% 112 1.2% 161 Ostrea or Crassostrea spp 10 0.2% 2 0.0% 20 0.4% 32 14 0.2% 4 0.0% 41 0.5% 59 Ostrea sp A (posible Ostrea conchapila 0.0% 0.0% 8 0.1% 8 0.0% 0.0% 10 0.1% 10 Saccostrea palmula 2 0.0% 3 0.1% 8 0.1% 13 3 0.0% 4 0.0% 20 0.2% 27 Cumingia sp 1 0.0% 0.0% 0.0% 1 1 0.0% 0.0% 0.0% 1 Cyclinella sp 0.0% 1 0.0% 1 0.0% 2 0.0% 1 0.0% 1 0.0% 2 Donax sp 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 2 0.0% 2 Dosinia dunkeri 3 0.1% 4 0.1% 31 0.6% 38 5 0.1% 8 0.1% 60 0.7% 73 Dosinia sp 4 0.1% 8 0.1% 85 1.6% 97 5 0.1% 14 0.2% 179 2.0% 198 Iphigenia altior 1 0.0% 3 0.1% 6 0.1% 10 1 0.0% 5 0.1% 8 0.1% 14 Isognomon sp 1 0.0% 0.0% 1 0.0% 2 1 0.0% 0.0% 1 0.0% 2 Leporimetis dombei 0.0% 1 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 Mactra fonsecana 1 0.0% 0.0% 0.0% 1 1 0.0% 0.0% 0.0% 1 Mactra sp 2 0.0% 3 0.1% 1 0.0% 6 4 0.0% 4 0.0% 1 0.0% 9 Mactra vanattae 1 0.0% 0.0% 0.0% 1 1 0.0% 0.0% 0.0% 1 Mactridae 0.0% 2 0.0% 7 0.1% 9 0.0% 6 0.1% 7 0.1% 13 Mulinia sp 0.0% 0.0% 11 0.2% 11 0.0% 0.0% 12 0.1% 12 Table 5.3: Bivalve assemblage at Vampiros-1

104

1 2a 2b 1 2a 2b Taxonomy MNI % MNI % MNI % TotalMNI NISP % NISP % NISP % TotalNISP Mytella (cf) guyanensis 0.0% 2 0.0% 3 0.1% 5 0.0% 2 0.0% 3 0.0% 5 Mytella spp 9 0.2% 15 0.3% 61 1.1% 85 15 0.2% 20 0.2% 65 0.7% 100 Pinctada mazatlanica 0.0% 0.0% 0.0% 0 0.0% 0.0% 0.0% 0 Pitar paytiensis 51 0.9% 10 0.2% 37 0.7% 98 100 1.1% 21 0.2% 70 0.8% 191 Pitar sp 21 0.4% 17 0.3% 129 2.4% 167 40 0.4% 30 0.3% 245 2.7% 315 Polymesoda boliviana 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 0.0% 1 Polymesoda sp 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 2 0.0% 2 Protothaca asperrima 74 1.4% 65 1.2% 22 0.4% 161 159 1.8% 121 1.3% 42 0.5% 322 Protothaca beili 1 0.0% 1 0.0% 3 0.1% 5 1 0.0% 1 0.0% 5 0.1% 7 Protothaca sp 71 1.3% 60 1.1% 87 1.6% 218 148 1.6% 122 1.4% 178 2.0% 448 Tagelus dombeii 1 0.0% 0.0% 0.0% 1 1 0.0% 0.0% 0.0% 1 Tagelus sp 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 0.0% 1 Tellina cf ecuadoriana 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 0.0% 1 Tellina laceridens 4 0.1% 3 0.1% 20 0.4% 27 10 0.1% 7 0.1% 37 0.4% 54 Tellina spp 3 0.1% 8 0.1% 87 1.6% 98 8 0.1% 22 0.2% 241 2.7% 271 Tivela cf argentina 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 2 0.0% 2 Tivela sp 0.0% 2 0.0% 1 0.0% 3 0.0% 2 0.0% 4 0.0% 6 Trachycardium procerum 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 0.0% 1 Trachycardium sp 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 0.0% 1 Unidentified bivalves 1 0.0% 0.0% 1 0.0% 2 1 0.0% 0.0% 1 0.0% 2 indt nacre 0.0% 0.0% 0.0% 0 0.0% 0.0% 0.0% 0

Table 5.3: Continued

105

1 2a 2b 1 2a 2b Taxonomy MNI % MNI % MNI % TotalMNI NISP % NISP % NISP % TotalNISP Calyptraea mamillaris 1 0.0% 0.0% 0.0% 1 1 0.0% 0.0% 0.0% 1 Cerithidea mazatlanica 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 0.0% 1 Cerithidea pulchra 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 0.0% 1 Cerithidea sp 0.0% 1 0.0% 3 0.1% 4 0.0% 1 0.0% 3 0.0% 4 Cerithidea valida 0.0% 0.0% 15 0.3% 15 0.0% 0.0% 15 0.2% 15 Potamididae 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 0.0% 1 Conus sp 1 0.0% 0.0% 0.0% 1 1 0.0% 0.0% 0.0% 1 Conus ximenes 1 0.0% 1 0.0% 2 0.0% 4 1 0.0% 1 0.0% 2 0.0% 4 Crepidula cf marginalis 1 0.0% 1 0.0% 1 0.0% 3 1 0.0% 1 0.0% 1 0.0% 3 Crepidula sp 2 0.0% 0.0% 2 0.0% 4 2 0.0% 0.0% 3 0.0% 5 Crucibulum sp 0.0% 0.0% 2 0.0% 2 0.0% 0.0% 3 0.0% 3 Cymatium sp 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 0.0% 1 Cymatium wiegmanni 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 0.0% 1 Littorina sp 5 0.1% 2 0.0% 1 0.0% 8 5 0.1% 2 0.0% 1 0.0% 8 Littorina varia 2 0.0% 6 0.1% 6 0.1% 14 2 0.0% 6 0.1% 6 0.1% 14 Malea ringens 1 0.0% 2 0.0% 12 0.2% 15 2 0.0% 3 0.0% 14 0.2% 19 Melampus carolianus 0.0% 0.0% 2 0.0% 2 0.0% 0.0% 2 0.0% 2 Melongena patula 2 0.0% 1 0.0% 1 0.0% 4 2 0.0% 1 0.0% 1 0.0% 4 Nassarius luteostomus 0.0% 2 0.0% 1 0.0% 3 0.0% 2 0.0% 1 0.0% 3 Nassarius sp 0.0% 1 0.0% 3 0.1% 4 0.0% 1 0.0% 4 0.0% 5 Nassarius versicolor 0.0% 0.0% 2 0.0% 2 0.0% 0.0% 2 0.0% 2 Natica sp 120 2.2% 271 5.0% 801 14.6% 1192 204 2.3% 395 4.4% 1177 13.0% 1776 Natica unifascita 286 5.2% 396 7.2% 803 14.7% 1485 451 5.0% 694 7.7% 1424 15.8% 2569 Naticidae 0.0% 0.0% 0.0% 0 0.0% 0.0% 0.0% 0 Oliva incrassata 0.0% 0.0% 3 0.1% 3 0.0% 0.0% 3 0.0% 3 Olivella sp 0.0% 0.0% 0.0% 0 0.0% 0.0% 0.0% 0 Olivella volutella 1 0.0% 0.0% 0.0% 1 1 0.0% 0.0% 0.0% 1 Pleuroploca granosa 3 0.1% 1 0.0% 16 0.3% 20 4 0.0% 1 0.0% 25 0.3% 30 Polinices cf otis 6 0.1% 4 0.1% 17 0.3% 27 6 0.1% 5 0.1% 20 0.2% 31 Polinices sp 1 0.0% 2 0.0% 8 0.1% 11 1 0.0% 2 0.0% 10 0.1% 13 Polinices uber 1 0.0% 8 0.1% 16 0.3% 25 1 0.0% 8 0.1% 16 0.2% 25 Prunum sapotilla 0.0% 1 0.0% 6 0.1% 7 0.0% 1 0.0% 6 0.1% 7 Prunum sp 0.0% 0.0% 0.0% 0 0.0% 0.0% 0.0% 0 Semicassis centiquadrata 1 0.0% 3 0.1% 1 0.0% 5 1 0.0% 3 0.0% 3 0.0% 7 Cassis sp 0.0% 1 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 Terebra cf robusta 0.0% 1 0.0% 1 0.0% 2 0.0% 1 0.0% 2 0.0% 3 Terebra sp 0.0% 0.0% 2 0.0% 2 0.0% 0.0% 2 0.0% 2 Thais kiosquiformis 132 2.4% 94 1.7% 299 5.5% 525 216 2.4% 167 1.8% 510 5.6% 893 Thais sp o Stramonita sp 1 0.0% 1 0.0% 2 0.0% 4 1 0.0% 2 0.0% 2 0.0% 5

Table 5.4: Gastropod assemblage at Vampiros-1

106

1 2a 2b 1 2a 2b Taxonomy MNI % MNI % MNI % TotalMNI NISP % NISP % NISP % TotalNISP Stramonita haemastoma 1 0.0% 2 0.0% 5 0.1% 8 1 0.0% 2 0.0% 9 0.1% 12 Indet. Gastropod 0.0% 3 0.1% 4 0.1% 7 0.0% 3 0.0% 6 0.1% 9 Indet. terrestrial Snail 1 0.0% 1 0.0% 0.0% 2 1 0.0% 2 0.0% 0.0% 3

Table 5.4: Continued

107

1 2a 2b 1 2a 2b Taxonomy MNI % MNI % MNI % TotalMNI NISP % NISP % NISP % TotalNISP Anadara cf formosa 0.0% 1 0.0% 0.0% 1 0.0% 2 0.1% 0.0% 2 Anadara similis 6 0.2% 1 0.0% 2 0.1% 9 7 0.2% 1 0.0% 2 0.1% 10 Anadara sp or Grandiarca sp 70 2.7% 22 0.9% 33 1.3% 125 70 1.9% 23 0.6% 34 0.9% 127 Anadara tuberculosa 72 2.8% 24 0.9% 16 0.6% 112 145 3.9% 43 1.2% 27 0.7% 215 Andara esmeralda 2 0.1% 0.0% 1 0.0% 3 2 0.1% 0.0% 1 0.0% 3 Grandiarca grandis 84 3.3% 26 1.0% 11 0.4% 121 168 4.5% 50 1.4% 18 0.5% 236 Argopecten sp 0.0% 0.0% 136 5.3% 136 0.0% 0.0% 136 3.7% 136 Argopecten ventricosa 0.0% 0.0% 89 3.5% 89 0.0% 0.0% 149 4.0% 149 Brachidontes sp 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 0.0% 1 Chione subrugosa 7 0.3% 2 0.1% 1 0.0% 10 17 0.5% 3 0.1% 1 0.0% 21 Cosmioconcha modesta 0.0% 1 0.0% 2 0.1% 3 0.0% 1 0.0% 2 0.1% 3 Crassostrea corteziensis 8 0.3% 0.0% 2 0.1% 10 13 0.4% 0.0% 2 0.1% 15 Ostrea or Crassostrea spp 1 0.0% 2 0.1% 12 0.5% 15 1 0.0% 2 0.1% 24 0.6% 27 Sacosstrea palmula 1 0.0% 1 0.0% 6 0.2% 8 1 0.0% 1 0.0% 13 0.4% 15 Donacidae 0.0% 0.0% 0.0% 0 0.0% 0.0% 0.0% 0 Dosinia dunkeri 5 0.2% 7 0.3% 3 0.1% 15 7 0.2% 11 0.3% 6 0.2% 24 Dosinia sp 5 0.2% 24 0.9% 34 1.3% 63 7 0.2% 48 1.3% 62 1.7% 117 Iphigenia altior 0.0% 0.0% 6 0.2% 6 0.0% 0.0% 15 0.4% 15 Mactra fonsecana 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 0.0% 1 Mactra sp 1 0.0% 0.0% 0.0% 1 3 0.1% 0.0% 0.0% 3 Mactridae 3 0.1% 0.0% 2 0.1% 5 3 0.1% 0.0% 3 0.1% 6 Mulinia sp 0.0% 0.0% 3 0.1% 3 0.0% 0.0% 5 0.1% 5 Mytella cf guyanensis 0.0% 1 0.0% 3 0.1% 4 0.0% 1 0.0% 6 0.2% 7 Mytella spp 31 1.2% 9 0.4% 68 2.7% 108 31 0.8% 9 0.2% 68 1.8% 108 Mytella strigata 0.0% 1 0.0% 3 0.1% 4 0.0% 1 0.0% 4 0.1% 5 Pitar paytiensis 47 1.8% 22 0.9% 42 1.6% 111 84 2.3% 43 1.2% 70 1.9% 197 Pitar sp 33 1.3% 19 0.7% 31 1.2% 83 62 1.7% 29 0.8% 55 1.5% 146 Polymesoda boliviana 3 0.1% 1 0.0% 0.0% 4 3 0.1% 1 0.0% 0.0% 4 Polymesoda sp 2 0.1% 0.0% 0.0% 2 3 0.1% 0.0% 0.0% 3

Table 5.5: Bivalve assemblage at Vampiros-2

108

1 2a 2b 1 2a 2b Taxonomy MNI % MNI % MNI % TotalMNI NISP % NISP % NISP % TotalNISP Prothotaca beili 0.0% 1 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 Protothaca asperrima 72 2.8% 14 0.5% 2 0.1% 88 138 3.7% 24 0.6% 3 0.1% 165 Protothaca sp 37 1.4% 8 0.3% 5 0.2% 50 72 1.9% 11 0.3% 8 0.2% 91 Pteria sterna 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 0.0% 1 Tellina laceridens 2 0.1% 4 0.2% 2 0.1% 8 3 0.1% 7 0.2% 3 0.1% 13 Tellina spp 1 0.0% 8 0.3% 15 0.6% 24 2 0.1% 15 0.4% 28 0.8% 45 Tivela sp 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 0.0% 1 unidentified bivalve 0.0% 1 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 Indt Nacre 0.0% 0.0% 0.0% 0 0.0% 0.0% 0.0% 0

Table 5.5: Continued

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1 2a 2b 1 2a 2b Taxonomy MNI % MNI % MNI % TotalMNI NISP % NISP % NISP % TotalNISP Calyptrea sp 1 0.0% 0.0% 0.0% 1 1 0.0% 0.0% 0.0% 1 Cerithidea mazatlanica 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 0.0% 1 Cerithidea sp 0.0% 0.0% 4 0.2% 4 0.0% 0.0% 4 0.1% 4 Cerithidea valida 0.0% 0.0% 29 1.1% 29 0.0% 0.0% 31 0.8% 31 Crepidula cf marginalis 1 0.0% 0.0% 0.0% 1 1 0.0% 0.0% 0.0% 1 Crepidula sp 0.0% 0.0% 7 0.3% 7 0.0% 0.0% 9 0.2% 9 Crucibulum sp 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 0.0% 1 Hexaplex sp 0.0% 0.0% 0.0% 0 0.0% 0.0% 0.0% 0 Littorina sp 2 0.1% 2 0.1% 1 0.0% 5 2 0.1% 2 0.1% 1 0.0% 5 Littorina varia 2 0.1% 2 0.1% 1 0.0% 5 2 0.1% 2 0.1% 1 0.0% 5 Malea ringens 0.0% 2 0.1% 1 0.0% 3 0.0% 2 0.1% 1 0.0% 3 Melongena patula 2 0.1% 0.0% 1 0.0% 3 2 0.1% 0.0% 2 0.1% 4 Nassarius luteostomus 1 0.0% 0.0% 2 0.1% 3 1 0.0% 0.0% 2 0.1% 3 Nassarius sp 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 2 0.1% 2 Natica sp 61 2.4% 51 2.0% 96 3.7% 208 99 2.7% 86 2.3% 163 4.4% 348 Natica unifasciata 215 8.4% 377 14.7% 309 12.0% 901 245 6.6% 402 10.9% 494 13.4% 1141 Naticidae 0.0% 0.0% 0.0% 0 0.0% 0.0% 0.0% 0 Pleuroploca granosa 0.0% 1 0.0% 2 0.1% 3 0.0% 1 0.0% 2 0.1% 3 Polinices cf otis 0.0% 6 0.2% 3 0.1% 9 0.0% 7 0.2% 3 0.1% 10 Polinices sp 1 0.0% 1 0.0% 0.0% 2 1 0.0% 1 0.0% 0.0% 2 Polinices uber 7 0.3% 16 0.6% 4 0.2% 27 7 0.2% 16 0.4% 4 0.1% 27 Prunum sapotilla 0.0% 1 0.0% 3 0.1% 4 0.0% 1 0.0% 3 0.1% 4 Prunum sp 1 0.0% 1 0.0% 0.0% 1 0.0% 1 Semicassis centriquadrata 1 0.0% 0.0% 0.0% 1 1 0.0% 0.0% 0.0% 1 Cassis sp 0.0% 1 0.0% 0.0% 1 0.0% 1 0.0% 0.0% 1 Stramonita haemastoma 0.0% 2 0.1% 4 0.2% 6 0.0% 3 0.1% 7 0.2% 10 Thais kiosquiformis 63 2.5% 28 1.1% 12 0.5% 103 104 2.8% 31 0.8% 18 0.5% 153 Indt gastropod 0.0% 4 0.2% 5 0.2% 9 0.0% 4 0.1% 5 0.1% 9 Indt landsnail 1 0.0% 0.0% 0.0% 1 1 0.0% 0.0% 0.0% 1

Table 5.6: Gastropod assemblage at Vampiros-2

Shellfish size and gathering methods

I measured the length and height of complete shells for the ten most abundant species to compare mollusc measurements between archaeological specimens at both shelters. The ten species are: Pacific calico scallop (Argopecten ventricosus), moon snail (Polinices cf otis), pitar venus clam (Pitar paytensis), single-banded moon snail (Natica unifasciata), grand ark (Grandiarca grandis), rock snail (Thais kiosquiformis), littleneck (Protothaca asperrima), black ark (Anadara tuberculosa), lacerate tellin (Tellina laceridens) and Dunker‘s dosinia (Dosinia dunkeri). I measured

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the length for bivalves and the height for gastropods using an electronic caliper.

Measurements are displayed in box plots. As a reference, I took length and height averages mentioned by Keen (1971).

I took these measurements to discern if change in mollusc size is related to gathering strategies. On the other hand, the comparison to modern size is to investigate if there is any change that could be related to human pressure over the resources or environmental change (Gassiot 2005; Cabral and Coelho 2003; Mannino and Thomas 2001;

Waselkov 1987).

Despite the small sample size, the box plots (Figures 5.6 to 5.15) show for all species, with the exception of the bivalves pitar venus clam and lacerate tellin, the average size of archaeological specimens in both shelters is smaller than modern ones described by Keen

(1971).

The samples of pacific calico scallop of both shelters (Figure 5.6) indicate that people were collecting specimens between 58.7 mm and 22.3 mm with a mean of 46.4mm in both

Vampiros-1 and Vampiros-2. Bigger animals were collected in Macrounit 2a than Macrounit

2b. The standard deviation of 4.3 mm for Vampiros-1 and 7.5mm for Vampiros-2 indicates that in the latter has greater size variation in the size of scallops collected than doesVampiros-1.

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Figure 5.6: Argopecten ventricosa´s boxplots. The line indicates Keen´s average length and then number enclosed on the box is the MNI.

In the case of moon snail (Figure 5.7), the specimens are between 44.2 to 15.7 mm in size with a mean of 28.9 mm inVampiros-1, and 25.3 mm in Vampiros-2. Their respective standard deviations (7.5 mm and 6.1 mm) indicate that there is higher size variation on

Vampiros-1 than in Vampiros-2. Big animals were collected in Macrounit 2b and Macrounit

1, and smaller animals in Macrounit 2a.

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Figure 5.7 : Polinices otis box plots. The line indicates Keen´s average height and then number enclosed on the box is the MNI.

For the pitar venus clam (Figure 5.8) the length ranges between 21 mm to 42.7mm in both shelters with a mean of 33.7mm at Vampiros-1 and 34.1 mm at Vampiros-2. Their correspondingly low standard deviations (3.6 mm and 3.7mm) suggest that there is no variation in shell size at either shelter. However, slightly smaller specimens were collected in

Macrounit 2b. In Macrounit 2a there is an increase in the length of the specimens, especially in Vampiros-2. In Macrounit 1 at Vampiros-1, specimens are either similar to Keen´s average specimens. In the same Macrounit at Vampiros-2, the specimens are bigger than

Keen´s average specimens.

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Figure 5.8: Pitar paytensis box plots. The line indicates Keen´s average length and then number enclosed on the box is the MNI.

The height of single banded moon snail samples (Figure 5.9) indicates that people were collecting specimens between 8.5 mm to 41.7mm in both shelters with a mean of

25.1mm at Vampiros-1 and 25.9 mm at Vampiros-2. The low standard deviation at the shelters (5.2 mm and 4.7 mm respectively) indicates that there is no variation in shell height in both shelters. The data show similar average´s sizes in each macrounit of both shelters.

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Figure 5.9: Natica unifasciata box plots. The line indicates Keen´s average height and then number enclosed on the box is the MNI.

The grand arks gathered by the Vampiros inhabitants (Figure 5.10) were smaller than the current average size. Larger animals were found in Macrounit 1 than in Macrounit 2a and

2b. The bivalves range between 10.5 mm to 91.4 mm in both shelters with a mean of 47.5 mm in Vampiros-1 and 46.2 mm on Vampiros-2. The standard deviations of Vampiros-

1(10.54 mm) and Vampiros-2 (11.2mm) indicate that there is size variation.

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Figure 5.10 : Grandiarca grandis box plots. The line indicates Keen´s average length and then number enclosed on the box is the MNI.

The height of the gastropod rock snail in the samples of Vampiros rockshelters

(Figure 5.11) indicates smaller animals were collected at the beginning of the occupation than at the end. The height ranges between 20.3mm to 49.1mm in both shelters with a mean of 34.6mm in Vampiros-1 and 35.6mm in Vampiros-2. The standard deviation of each shelter (5.1mm for Vampiros-1 and 5.2 mm for Vampiros-2) suggests that there is no variation of gastropod size

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Figure 5.11: Thais kiosquiformis box plots. The line indicates Keen´s average height and then number enclosed on the box is the MNI.

The littleneck (Figure 5.12) length ranges between 18.3 mm to 45.8 mm in both shelters with a mean of 30.1mm at Vampiros-1 and 32.3mm at Vampiros-2. The standard deviation of each shelter 4.1 mm and 3.9 mm indicates that there is no variation in shell size.

Smaller specimens than the Keen´s average were gathered by Vampiros inhabitants in

Macrounit 2a and Macrounit 2b than in Macrounit 1.

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Figure 5.12: Protothaca asperrima box plots. The line indicates Keen´s average length and then number enclosed on the box is the MNI.

Smaller specimens of black ark (Figure 5.13) were collected in Macrounit 2b than in

Macrounits 2a and 1. The specimens´ length ranges between 23.5 mm to 46.8 mm in both shelters with a mean of 32.1 mm in Vampiros-1 and 31.8 mm in Vampiros-2. The standard deviation is 4.4 mm at Vampiros-1 and 6.5 mm at Vampiros-2 suggesting that there is no variation in size in both shelters.

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Figure 5.13: Anadara tuberculosa box plots. The line indicates Keen´s average length and then number enclosed on the box is the MNI.

Specimens of lacerate tellin clams (Figure 5.14) larger than Keen´s average specimen were found in both shelters and all macrounits with the exception of those found at

Vampiros-1 in Macrounit 2b, which are smaller. The shell length varies between 26.1mm to

41.8 mm in both shelters with a mean of 34.2 mm at Vampiros-1 and 37.4mm at Vampiros-

2. Their standard deviation of 3.2 mm and 1.4 mm respectively indicating that there is no variation at either shelter.

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Figure 5.14: Tellina laceridens box plots. The line indicates Keen´s average length and then number enclosed on the box is the MNI.

Finally, the Dunker‘s dosinia clams (Figure 5.15) were collected at the two shelters in different ways. In Vampiros-2 smaller animals were collected in Macrounit 2a and 2b.

Vampiros-1 does not show a clear pattern, presenting similar lengths at Macrounits 2a and

2b. The shell length differs between 32.6 mm to 53.6 mm with a mean of 43.02 mm at

Vampiros-1 and 39.7 mm at Vampiros-2. The standard deviation of 4.1 mm and 10.9 mm, which indicates that there is bigger variation at Vampiros-2 than Vampiros-1

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Figure 5.15: Dosinia dunkeri box plots. The line indicates Keen´s average length and then number enclosed on the box is the MNI.

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State of Preservation, Human and Natural Modifications

Visual examination of molluscan remains indicates that natural and cultural factors affected their preservation. Taphonomic details offer some hints of their provenience as well.For each taxon per stratigraphic unit, I recorded the following characteristics by weight, e.g; completeness, encrustation, perforation, abrasion, acid dissolution, and heat treatment.

The inhabitants of both shelters gathered well-preserved shells. Most of the shells maintained their colors. There are few specimens where dead animals were gathered (Figure

5.16). The removal of calcium carbonate by near-shore waves, bioeroders or abrasion was identified in few cases at Vampiros, the most frequent being in Macrounit 1 (subunit 5),

Macrounit 2a (subunit 7) and Macrounit 2b (subunit 16, 22 and 29) at Vampiros-1; this modifications were observed at Vampiros-2 in Macrounit 1(subunit 6), Macrounit 2a

(subunit 28) and Macrounit 2b (subunit 43).

The dissolution of calcium carbonate by the presence of water or bioturbation (acid dissolution) was recognized at Vampiros. A chalky appearance was commonly observed in

Vampiros-1 in Macrounit 1(all subunits), Macrounit 2a (subunits 6 and 7) and Macrounit 2b

(subunits 15, 19, 20, and 29). For Vampiros-2 this characteristic was frequent in Macrounit

1(subunit 6), Macrounit 2a (subunits 8, 9, 23) and Macrounit 2b (subunits 33, 45) (Figure

5.17).

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Figure 5.16: Mollusk remains with abrasion per subunit in each macrounit and shelter by weight (gr). 123

Figure 5.17: Mollusk remains with acid dissolution per subunit in each macrounit and shelter by weight(gr).

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Figure 5.18: Mollusk remains with encrustation per subunit in each macrounit and shelter by weight (gr). 125

Encrustation or the growth of other organisms occurs on the surface of shells

(Figure 5.18). This process was observed at Vampiros in a few specimens, mostly on pacific calico scallop, made by barnacles and worms. In Vampiros-1 specimens with encrusted surfaces were observed in Macrounit 1 (subunits 3, 4, 5), Macrounit 2a (subunit 6) and

Macrounit 2b (subunits 24, 25, 26, 27, 28 and 29). In Vampiros-2 encrusted specimens were frequently identified in Macrounit 1(subunits 3, 6 and 7) Macrounit 2a (subunit 19) and

Macrounit 2b (subunits 46, 47, 48, 49, 51, 52, 53, 54, 55, 56, 57, 58 and 59).

Animal and human activities modified some shell specimens. In Vampiros-1 (see

Figures 5.19 and 5.20) there were frequent examples of complete specimens in Macrounit 1

(subunit 2 and 4) and lower frequency of fragmented and semi-fragmented specimens. In

Macrounit 2a (subunit 7 and hoyo 10) and Macrounit 2b fragmented specimens of both bivalves and gastropods were frequent. In Vampiros-2 complete specimens of gastropods and bivalves were observed in Macrounit 1 (subunit 7), Macrounit 2a (subunit 12, 13, 18, 19 and 27) and Macrounit 2b (subunits 32, 35 and 49).

Shell remains at Vampiros rockshelters were exposed inconsistently to heat (Figure

5.21). As observed in Vampiros-1 differential exposure to heat was observed from scorched to calcined shells at Macrounit 1(subunit 5), Macrounit 2a (subunit 6, 7 and 8) and Macrounit

2b (subunits 22, 25 and 28). Several degrees of heat exposure (e.g. from scorched to calcinated shells) were recognized at Vampiros-2 in Macrounit 1(subunit 3, 4 and 7),

Macrounit 2a (subunit 13 and hoyo 18) and Macrounit 2b (subunit 38, 39, 41 and hoyo 28).

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Figure 5.19: Bivalve remains completeness (fragment, semi-fragment, whole) per subunit in each macrounit and shelter by weight (gr). 127

Figure 5.20: Gastropod remains completeness (fragment, semi-fragment, whole) per subunit in each macrounit and shelter by weight(gr). 128

Figure 5.21: Mollusc remains heat treatment (black, grey, brown) per subunit in each macrounit and shelter by weight (gr). 129

Figure 5.22: Paired molluscan remains per subunit in each macrounit and shelter by weight(gr). 130

Figure 5.23: Mollusc remains with natural perforation per subunit in each macrounit and

shelter by weight(gr).

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Articulated bivalves although were rare overall (Figure 5.22). They were frequently recorded in Macrounit 2a (subunit 6) and Macrounit 2b (subunits 13, 23) in Vampiros-1. In

Vampiros-2 they were commonly observed in Macrounit 1 (subunit 3), and Macrounit 2b

(subunit 51).

Perforations caused by worms, barnacles, gastropods or bivalves were identified at

Vampiros. As shown in Figure (5.23), there were examples at Vampiros-1 in Macrounit 2a

(subunit 6) and Macrounit 2b (subunits 11, 20) and Macrounit 1 (subunit 3) and Macrounit

2b (subunits 36, 39) at Vampiros-2.

Vampiros-1

T rachycardium sp T ivela cf argentina T erebra sp T ellina laceridens T agelus dom beii Saccostrea palm ula Protothaca sp Polymesoda sp Polinices sp Pitar sp Ostrea sp A (posible Ostrea conchapila Olivella sp Natica sp Nassarius luteostomus M ulinia sp M alea ringens M actra sp Littorina sp

taxonomy Iphigenia altior Dosinia dunkeri Cymatium sp Crucibulum sp Crassostrea corteziensis Conus sp Chione obliterata Cerithidea pulchra Carditam era radiata Argopecten ventricosa Anadara tuberculosa Anadara perlabiata 0.00 0.50 1.00 1.50 Completeness Index

Figure 5.24: Completeness ranges among taxa at Vampiros-1.

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Vampiros- 2

Tellina spp Sem icassis centriquadrata Prunum sp Protothaca asperrim a Polymesoda boliviana Polinices cf otis Pitar paytiensis Natica unifasciata Nassarius luteostomus Mytella cf guyanensis M alea ringens M actra fonsecana

Taxonomy Iphigenia altior Dosinia sp Crucibulum sp Crassostrea corteziensis Cerithidea valida Cassis sp Argopecten ventricosa Anadara tuberculosa Anadara cf formosa 0.0 1.0 2.0 3.0 Completeness Index

Figure 5.25: Completeness ranges among taxa at Vampiros-2.

To observe if completeness varied between taxa, I plotted the completeness index

(MNI/NISP) versus species. As can be observed, in Vampiro-1 (Figure 5.24) black ark

(Anadara tuberculosa) was the most fragmented species followed by single/banded moon snail

(Natica unifasciata), grand ark (Grandiarca grandis), rock snail (Thais kiosquiformis) and littleneck

(Protothaca asperrima). But at Vampiros-2 data suggest that little neck (Protothaca asperrima) was the most fragmented species followed by rock snail (Thais kiosquiformis), ark (Anadara sp),

Dunker‘s dosinia (Dosinia dunkeri), mussel (Mytella sp), single banded moon snail (Natica unifasciata ) and pitar venus clam (Pitar paytensis) ( Figure 5.25).

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Figure 5.26: Heat treatment among taxa at Vampiros-1.

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Taxa Vampiros-2

Tellina spp Stramonita haemastoma Prunum sp Protothaca asperrima Polymesoda boliviana Polinices sp Pleuroploca granosa Pitar paytiensis Natica unifasciata Nassarius sp Mytella spp Malea ringens Iphigenia altior Dosinia sp Crepidula sp Cosmioconcha modesta Cerithidea valida Cassis sp Argopecten sp Anadara sp or Grandiarca sp

0.0 200.0 400.0 600.0

Total burnt

Figure 5.27: Heat treatment among taxa at Vampiros-2.

To observe if heating varied among taxonomic groups, I plotted the total weight of burned species. For Vampiros-1 single-banded moon snail was the species most affected by heating, followed by rock snail, grand ark, pitar venus clam, and littleneck (Figure 5.26). For

Vampiros-2 ark (Anadara sp) was the most affected by heating followed by Dunker‘s dosinia, single-banded moon snail, pitar venus clam, littleneck and rock snail (Figure 5.27).

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Interpretation

As indicated by relative frequencies, marine shellfish exploitation in Vampiros was based mainly on the gastropods rock snail, moon snail and single-banded moon snail and the bivalves littleneck, pitar venus clam, gran ark, mussel , lacerate tellin , black ark, Dunker´s dosinia and Pacific calico scallop . Shellfish gathering focused on intertidal mollusc‘s i. e, those associated with the nearest environments to the site. This environment of Pre-

Columbian exploitation focused on the low intertidal zone, mudflats, shallow waters, firm substrates and gravel or sand in strong currents substrate. The subunits corresponding to the

Macrounits 2a and 2b in both shelters are dominated by firm intertidal substrate species such as Cortez oyster , little neck and pitar venus clam, low intertidal zone species such as the tall false donax (Iphigenia altior), Dunker´s dosinia , single-banded moon snail and to a lesser extent in Macrounit 1, mangrove species such as black ark , rock snail and pacific melongena (Melongena patula), intertidal mud species such as mussel, horn shell (Cerithidea sp), pacific cask shell (Malea ringens), moon snail , granose horse conch (Pleuroploca granosa) , tellin (Tellina sp), olivella snail (Olivella sp), marginella (Prunum sp), auger (Terebra sp) and sandbars at extreme low tide species such as grand ark (Grandiarca grandis).

The mollusc data indicate that at the beginning of occupation at both shelters

(Macrounit 2b), gravel or sand in strong currents substrate and clear water environments were available for the pre-Columbian inhabitants to exploit species such as Pacific calico scallop (Carvajal Contreras, et al. 2008).

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As mentioned in chapter 2, there is no detailed geomorphologic information that explains the position of Cerro Tigre on the ancient coastline and the changes that occurred to the Santa Maria River; however the low frequency of brackish water species such as marsh clam (Polymesoda boliviana) or sandy species such as Panamic bonnet (Cassis centiquadrata),

Donax sp and Tivela sp. implies that the prograding coastline and delta formation had not completely developed at Cerro El Tigre and surroundings.

The diversity indices indicate utilization of several mollusc species and consequently that native people were focusing their shell gathering toward several taxa. Almost all the 54 species were presented in the same proportion in each Macrounit in both shelters and one taxon did not dominate each Macrounit.

The top ranked species collected for pre-Columbian inhabitants, perhaps as an occasional meal, were from the gastropods single-banded moon snail (Natica unisfasciata), moon snails( Polinices uber and Polinices cf otis); and rock snail, and from the bivalves the littleneck, black ark, grand ark, pitar venus clam, Dunker‘s dosinia, lacerate tellin, Cortez oyster, mussel and pacific calico scallop .

Less frequent taxa such as the gastropods the Wiegmann‘s triton (Cymatium wiegmanni), Nassa (Nasarius sp), dovesnail (Cosmioconcha modesta) ,circular Chinese hat

(Calyptrea sp), periwinkle (Littorina sp), pacific melongena (Melongena patula), cone snail (Conus sp), crepidule (Crepidula sp), melampus (Melampus carolianus), marginella (Prunum sp) and

Olivella snail (Olivella sp) and the bivalves tagelus (Tagelus sp), , tall false donax (Iphigenia altior), cardita (Carditamera sp), surfclam(Mactra sp) and chione (Chione sp) could reflect accidental collection. These species share the same habitat as the main species collected by

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Pre-Columbian inhabitants or are part of the trophic system. For example, the Wiegmann‘s triton preys on the pacific calico scallop. Carnivorous snails such as moon snails and single- banded moon snails , rock snails ), cone snail, Olivella snail , pacific melongena, and auger feed on clams such as Chione sp, Dosinia, Donax, Tivela, Tellina and Tagelus (Cruz-Soto and

Jiménez 1994).

People collected these species at different stages of their growth, although as the comparison of average size between archaeological mollusc and modern ones indicate that small sizes or juvenile specimens were frequent, especially in the case of grand ark. Other plausible explanation of the small size of these intertidal molluscs may well have resulted in overexploitation and stock collapses.

Expedient tools and little effort would have been used to collect shellfish, with the exception of Pacific calico scallop. The single-banded moon snail is easily obtained by hand during the ebb tide. Digging tools would have been employed to remove Chione and Tivela.

Stronger tools such as stone axes or choppers would have been applied to unfasten periwinkle, black ark, Cortez oyster, horn shell and rock snails from the mangroves. The heterogeneity in size and accidental gathering of some species, as mentioned above, suggests a mass collection or unselective procurement method (Carvajal Contreras 1998). I said the

Pacific calico scallop is an exception because today, this scallop is obtained in the Pearl islands using an artisanal trawling method attached to small traditional boats in shallow areas

(Medina, et al. 2007), but it also can be easily obtained during extreme tides called aguajes, which generally occur during the dry season associated with el Niño events (Cooke, personal communication; Rodriguez, personal communication).

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Finally, my assumption that the apparent decrease in the sizes of shells of single- banded moon snail from the bottom of Macrounit 2 to the top of Macrounit 1 is not supported by the data (Carvajal Contreras, et al. 2008). The single-banded moon snail displays small size throughout the entire stratigraphic sequence.

Natural processes and the activities of the peoples who occupied the shelters have had a profound post-depositional impact on the integrity and preservation of shell remains.

Some of these agents left a signature on shells. For example, erosion of the shell surfaces owing to a differential sediment movement was commonly found on shells collected from the beaches such as the grand ark, single-banded moon snail, pitar venus clam, , Pacific calico scallop and Dunker‘s dosinia (Claassen 1998; Rick, et al. 2006). Most of these specimens were found in Vampiros-1 in ashy layers.

In general, shell specimens are soon disarticulated after death; therefore articulated shells indicate a rapid burial. This feature was observed in the pitar venus clam, lacerate tellin, grand ark, littleneck and black ark in organic sediments and compact ashy layers in both shelters in all Macrounits (Claassen 1998; Rick, et al. 2006).

Acid dissolution of shell carbonate represents fluctuations in temperature, pH or carbon dioxide in surrounding waters (Claassen 1998; Rick, et al. 2006). This feature was observed in almost all the species in the top layers of Vampiros-1 and Vampiros-2.

Barnacles, serpulid worms and other organisms grew on the surfaces of single-banded moon snail, pitar venus clam, oyster, mussel and tall false donax and especially pacific calico scallop. Perhaps these shells were exposed above the sediment- water interface as was

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observed in the bottom layers of Vampiros-1 and Vampiros-2(Claassen 1998; Rick, et al.

2006).

Animal burrowing and human activities (i.e. trampling, digging postholes, cooking) would have been a factor contributed to the fragmentation of shells (Claassen 1998; Rick, et al. 2006). In both shelters complete specimens were found at the top (Macrounit 1), whereas semi-fragmented and fragmented specimens were common in Macrounit 2 in association with anthropogenic features such as postholes ash layers and hearths.

Shells range from light grey to black and scorched-brown. It is not possible to determine if these colors are related to cooking or are post-depositional effects from the hearths used to preserve fish (Claassen 1998; Rick, et al. 2006). Fragmentation would have occurred from a combination of heating and trampling. Distinct processing treatments or post-depositional events such as fragmentation and heat alteration are apparent in

Vampiros-1 and Vampiros-2 for the single-banded moon snail, black ark, littleneck and rock snail. The trampling affected fragile species such as surfclam, mussel and tellin. The differences in mollusk´s fragmentation between Vampiros-1 and Vampiros-2 would have also be related to pre-Columbian inhabitants lived at Vampiros-2 and processed fish at

Vampiros-1.

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Chapter 6

Other vertebrate remains at Cueva de los Vampiros Site (AG-145)

Introduction

In this chapter, I focus on the non- fish vertebrate faunal remains. The faunal assemblage was recovered from excavations conducted between 2002 and 2006. In 2002-2004, these pits were excavated using arbitrary 5 to 20 centimeter levels, and the excavated material was screened through 1/8‖ mesh. Samples from 2005 were recovered using 1/16‖ mesh.

Numerous diagnostic fragments of mammals, amphibians, reptiles and birds were present in the deposits.

Analytical procedures

As described in chapter 1, Cueva de los Vampiros deposits were divided into three analytical units. Macro unit 1 is a soft soil that contains pottery, stone tools and faunal remains.

Macrounit 2 is a complex deposit containing hearths, ashes, post-molds, pottery, lithic, shell and vertebrate remains, and was subdivided into macrounit 2a and macrounit 2b.

Each bone in Vampiros´ samples was identified through anatomical landmarks to the lowest taxonomic level possible using the comparative vertebrate skeleton collection housed at the Smithsonian Tropical Research Institute in Panama. Field guide and identification manuals were also consulted (Olsen 1968, 1979; Wyneken 2001). Some of the bones were analyzed by Maximo Jiménez. The bones were sorted by class and identified taxonomically.

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Later, the skeletal element, side, portion, weight and taphonomic features such as fragmentation, burning, cut marks or other modifications were recorded (Blasco 2008;

Lyman 1994; Reitz and Wing 1999a).

Cueva de los Vampiros vertebrate archeofauna were quantified using number of identified specimens (NISP) that included both complete and fragmented bones for each specific taxon (Ringrose 1993). The minimum number of individuals (MNI) was determined as well for each taxonomic group, taking into account the most abundant anatomical element and size ranges (Ringrose 1993). I estimated specimen size in grams using the proportional method (Cooke, et al. 2004: 215).

Results The Cueva de los Vampiros non-fish vertebrate fauna reported here consists of 541 bone specimens identified to class level as illustrated in Table 6.1. Although it is a small sample, further excavations could increase its diversity and proportions. Reptiles dominated (60%), followed by mammals (27%), amphibians (7%) and birds (6%). Twenty- five genera and 18 species representing 25 families are identified.

Macrounit Amphibia NISP Aves NISP Mammalia NISP% Reptilia NISP (NISP) % (NISP) % (NISP) (NISP) %

1 36 0.07 29 0.05 109 0.2 229 0.42

2a 0 0 4 0.008 24 0.04 63 0.12

2b 0 0 1 0.002 15 0.003 32 0.06

Total 36 0.07 34 0.06 148 0.27 324 0.6

Table 6.1: Non- fish vertebrate class frequencies from Vampiros-1 and Vampiros- 2 expressed in number of identifiable specimens (NISP and NISP %) 142

The identified vertebrate fauna (Table 6.2) in comparison with fish demonstrated that bony fish are the dominant class at Vampiros rockshelters (NISP 96%), followed by other vertebrates (NISP= 3.8%) and then by cartilaginous fish (NISP= 0.2%). These general trends hold for all three macrounits. In the next sections, each macrounit is described in detail.

Macrounit Chondrichthyes NISP Osteichthyes NISP % Other NISP Total (NISP) % (NISP) Vertebrat % es (NISP) 1 8 0.06 2151 15.3 403 2.8 2562 2a 7 0.05 6119 43.5 91 0.6 6217 2b 13 0.09 5233 37.2 48 0.4 5294 Total 28 0.2 13503 96 542 3.8 14073

Table 6.2: Fish versus Non- fish vertebrate from Vampiros-1 and Vampiros- 2 expressed in number of identifiable specimens (NISP and NISP %)

Macrounit 1

Reptiles are the first most common non-fish vertebrate represented in this unit (Table 6.3).

This deposit includes freshwater turtles (Kinosternon and Trachemys scripta), sea turtles

(Eretmochelys imbricate, Chelonia agassizii), lizards (Ctenosaura similis and Iguana iguana), and snakes

(Boa constrictor).

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Reptiles Specie NISP MNI Biomass Boa constrictor 44 4 6850 cf Iguanidae 1 cf Trachemys 1 Chelonia agassizii 26 9 377000 Cheloniidae 49 Ctenosaura similis 46 18 9750 Eretmochelys imbricata 2 1 8000 Iguana iguana 38 18 25200 Iguanidae 7 Kinosternon 3 1 300 Lacertilia 8 3 Serpentes 1 Testudines 1 Trachemys scripta 2

Table 6.3: Reptiles at macrounit 1(NISP, MNI and estimated biomass)

The freshwater turtles are represented by bones from the axial skeleton (plastron, peripheral bones and costal scutes) and a fragment of a pelvis. Some of these costal scutes show evidence of burning. Nowadays, Kinosternon and Trachemys scripta are available in the vicinities of Cueva de los Vampiros.

Figure 6.1: Element distribution of freshwater turtle expressed in number of identified specimens (NISP) 144

Macrounit 1

No Bone NISP modification Type Location Burning Action Freshwater turtles (Trachemys & black and cooking? Kinosternon) costal 3 scorched smoking? irregular missing distal disarticulation coracoid 1 1 fracture portion ? irregular missing distal disarticulation quadrate 1 1 fracture portion ? irregular missing distal disarticulation ulna 1 1 fracture portion ? present irregular lateral and disarticulation dentary 2 2 fracture distal portion ? irregular missing distal disarticulation metacarpo 1 1 fracture portion ? irregular peripheal bone 11 11 fracture cranial- caudal defleshing? missing irregular proximal cooking? pterygoid 1 1 fracture portion smoking? detached irregular vertebrae cooking? vertebrae 12 12 fracture epiphysis smoking? irregular flat bone 4 4 fracture defleshing? Marine turtles irregular cooking? (Cheloniidae) shaft 1 1 fracture smoking?

irregular almost cooking? Exoccipital 1 1 fracture complete smoking? irregular missing distal cooking? rib 1 1 fracture portion smoking? irregular cooking? femur 2 2 fracture shaft smoking? 1 shaft, 1 irregular missing distal disarticulation fibula 3 2 fracture portion ? cooking? humerus 2 smoking? irregular cooking? long bone 1 1 fracture shaft smoking? 2 shafts, 1 irregular missing distal cooking? metapodial 3 3 fracture portion smoking? cooking? premaxilla 1 smoking? proximal shaft irregular and detached cooking? radius 1 1 fracture epiphysis smoking? irregular cooking? tibia 1 1 fracture shaft smoking? 1 detached irregular vertebrae disarticulation Lizards vertebrae 6 1 fracture epiphysis ? (Ctenosaura & irregular missing distal disarticulation Iguana) jugal 1 1 fracture portion ?

Table 6.4: Reptiles at macrounit 1: modifications (NISP)

Large marine sea reptiles correspond to Chelonia agassizii and Eretmochelys imbricata in this unit (Table 6.4 and Figure 6.1). Fragmented bones with evidence of oblique or transverse irregular breaks (Reitz and Wing 2008: Figure 6.6, 169 )and burning marks are 145

distributed in axial skeleton (vertebrae, peripheral bones), forelimbs (radius, metacarpal), hindlimbs (fibula, tibia, and femur), palatal area (pterygoid, quadrate), pectoral girdle

(coracoids) and the jaws (dentary).

Figure 6.2: Element distribution of marine turtles expressed in number of identified specimens (NISP) Ctenosaura similis and Iguana iguana are represented by bones from the axial skeleton

(vertebrae, ribs), forelimbs (humerus, ulna, radius, and phalanges), hindlimbs (fibula, tibia, and femur), skull (basisphenoides, exoccipital, frontal and basioccipital), pelvis, platal area

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(jugal) and jaws (dentary) (Figure 6.3). In the sample there are juvenile individuals, fragmented bones with evidence of irregular fracture and burning.

Figure 6.3: Element distribution of Iguana expressed in number of identified specimens (NISP) Finally, vertebrae and a dentary corresponding to Boa constrictor were identified. These bones do not have evidence of being burned or butchered.

The mammals are the second class in importance and are represented by small rodents (Oryzomys, Lyomys adspersus, and Sigmodon hispidus), medium-size mammals (Caluromys,

Procyon lotor, and Cuniculus paca), ungulates (Odocoileus virginianus), bats (Chiroptera) and human remains (Table 6.5).

Few bones from the skull and limbs identified as small rodents were found in this assemblage and do not present evidence of cut or burn marks. These bones could represent

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intrusions since these rodents use burrows dug by other animals as nesting places. Bat remains could also be interpreted as intrusions in this deposit since today hematophagous species live in the shelters (Reid 1997). Grouped remains of at least 4 species of these animals were found with no evidence of being used as food (Carvajal Contreras, et al. 2008).

Mammalia Specie NISP MNI Biomass Artyodactyla 1 1 Caluromys 1 cf Cricetidae 1 cf Rodentia 1 Chiroptera 65 20 Cuniculus paca 1 1 Didelphidae 1 Didelphis marsupialis 1 Homo sapiens 5 Lyomis adspersus 2 1 50 Mammalia 2 Muridae 1 1 40 Odocoileus virginianus 4 2 Oryzomys 1 1 Procyon cf lotor 22 14 1450 Sigmodon hispidus 2 1

Table 6.5: Mammals macrounit 1 (NISP, MNI and estimated biomass)

Medium-size mammals such as raccoon, opossum and paca were identified from macrounit 1 (Table 6.6). These remains present evidence of being burned and correspond to the axial skeleton (ribs), limbs (humerus, scapula femur, tibia, radius, phalanges and metatarsals), pelvis and jaws (dentary) (Figures 6.4, 6.5 and 6.6). The raccoon is the most common medium-size mammal in the sample. Some of these bones are juvenile specimens.

One radius exhibits a uniform cut mark. 148

Figure 6.4: Element distribution of small mammals expressed in number of identified specimens (NISP)

Bone fragments corresponding to at least two deers were found at macrounit 1.

These are unburned bone from hind limbs (tibia with postdepositional fracture), carpals

(unciform and magnum) and fragmented ribs.

Finally, heavily fragmented human remains were identified at Cueva de los Vampiros represented by a cranium and one tooth. These bones could have been the result of post - depositional burials.

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Table 6.6: Mammals at macrounit 1: modifications (NISP)

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Figure 6.5: Mammal and reptile bones and teeth from the Vampiros shelters, Cerro Tigre, Cocle, Panama (Photos: Tara Hornung). (a) Black iguana (Ctenosaura sp.), anterior right dentary (split longitudinally), (b) snake (cf. Boa constrictor), right dentary, Vampiros-2, (c) snake (cf. Boa constrictor), vertebra, probably same individual as (b), Vampiros-2, (d) hispid cotton rat (Sigmodon hispidus), right femur, distal epiphysis absent (unfused), Vampiros-1, (e) hispid cotton rat (Sigmodon hispidus), left mandible, probably same individual as (c), Vampiros- 1, (g) opossum (Didelphidae, probably Didelphis marsupialis), caudal vertebra, Vampiros-1, surface, (f) mud turtle (Kinosternon scorpioides), left ilium, Vampiros-2, (h) bat (Chiroptera), right humerus, proximal epiphysis absent (unfused), Vampiros-2, (j) bat (Chiroptera), unidentified species, left humerus, Vampiros-1, (k) bat (Chiroptera), left mandible, Vampiros-1 and (l) grayfox (Urocyon cinereoargenteus), left mandibular premolar.

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Figure 6.6: Mammal, bird and reptile bones from the Vampiros shelters, Cerro Tigre, Cocle´ , Panama (Photos: Tara Hornung). (a) Raccoon (Procyon cf lotor), right tibia,Vampiros-2, (b) paca (Agouti paca), distal left humerus, Vampiros-2, (c) anteater (Tamandua cf mexicana), proximal left ulna,- Vampiros-2, (d) black vulture (Coragyps atratus), right articular, Vampiros-1, (e) brown pelican (Pelecanus occidentalis), proximal left radius,Vampiros-2, (f) cormorant (Phalacrocorax cf olivaceus), left coracoid, Vampiros-2; (g) green iguana (Iguana iguana), right dentary, Vampiros-1, (h) black iguana (Ctenosaura sp.), frontal, Vampiros-1, (j) green iguana (Iguana iguana), left humerus, Vampiros-2 and (k) green turtle (Chelonia agassizzi), anterior dentary, Vampiros-1.

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Marine frog bones (Bufo marinus) were identified in this unit corresponding to at least 5 individuals (Table 6.7 and Figure 6.7). Although these animals have been used for food in pre-Columbian times (Cooke 1989), these bones do not present marks of having been burned or cut (Table 6.6). The bones of one of these individuals were found clustered, which suggests that this animal died in situ. These remains were intrusions in this deposit

(Carvajal Contreras, et al. 2008)

Amphibia Specie NISP MNI Biomass Bufos marinus 36 5 1280

Table 6.7: Amphibian at macrounit 1 (NISP, MNI and estimated biomass)

Table 6.8: Amphibian at macrounit 1: modifications (NISP)

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Figure 6.7: Partial skeleton of a marine toad (Bufo marinus), Vampiros-1.Maximum acetabular width (*) predicts an estimated body mass of 408 g, and a snout-vent length of 166mm (Cooke, 1989) (Photo: R.G. Cooke).CL—clavicle, CO—coracoid, DN—dentary, ET— ethmoid, EX—exoccipital,FE—femur, FR—frontal, H—humerus, IL—ilium, MA-maxilla, MT—metatarsal, PS—parasphenoid, PT—pterygoid, SU—suprascapular, TF—tibio-fibula, U—urostyle and V—vertebra ( from Carvajal Contreras, et al. 2008: 99)

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Birds are scarcely represented at macrounit 1 (Table 6.9 and Figure 6.6). Most of the bones are limb bones (humerus, tibiotarsus, coracoid, scapula, radius, phalanges) and lower jaw (dentary). Some of them come from immature specimens and there is no evidence of burn marks but fractures due postdepositional events (Table 6.10). One of these bird´s bones (Strigidae) has evidence of being worked as an adornment.

Aves Specie NISP MNI Biomass Aves 11 2 cf Aves 1 1 cf Cairina 1 1 cf Thraupinae 1 1 10 Coragyps atratus 2 2 Cuculidae 1 1 Laridae 2 2 100 Passeriformes 5 2 20 Phalacrocorax cf 1 3 700 punctatus olivieri Strigidae 1 1 350 Sturnella 1 1 160

Table 6.9: Birds at macrounit 1 (NISP, MNI and estimated biomass)

No Bone NISP modification Type Location Burning Action missing irregular distal coracoid 1 1 fracture portion postdepositional missing irregular distal dentary 1 1 fracture portion postdepositional 1 shaft, 1 missing distal portion, 1 missing 2 irregular proximal humerus 3 3 fracture portion bead missing irregular distal Birds radius 1 1 fracture portion disarticulation Table 6.10: Birds at macrounit 1: modifications (NISP)

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The species identified are a juvenile mottled owl (Ciccaba virgata), cormorant

(Phalacrocorax punctatus olivieri), owl (Strigidae), passerine birds (Cuculidae), gulls (Laridae),

Sturnella (Meadowlark) and Black vulture (Coragyps atratus). This last specimen and the owls are considered a product of post-depositional events since there is no evidence of modification in these bones. Black vulture and Owl bones presence at Vampiros rockshekters could be related to their dietary (it feeds at garbage dumps) or reproductive (it lays its eggs on the ground) behavior (Carvajal Contreras, et al. 2008; Ridgely and Gwynne

1989).

Macrounit 2a

Reptiles again are the most common class in this unit, followed by mammals and birds.

Species identified were the freshwater turtle, identified as the common slider (Trachemys scripta), lizards (Ctenosaura similis and Iguana iguana), and sea turtles (Chelonia agassizii and

Lepidochelys) (Table 6.11).

Reptiles Specie NISP MNI Biomass cf Cheloniidae 1 cf Iguanidae 1 Chelonia agassizii 13 9 555000 Cheloniidae 29 2 34000 Ctenosaura similis 6 5 2850 Iguana iguana 10 6 9250 Iguanidae 1 Lepydochelys 1 1 80000 Trachemys scripta 1 1 12000

Table 6.11: Reptiles at macrounit 2a (NISP, MNI and estimated biomass) 156

The common slider is represented by an unburned hypoplastron presenting an irregular fracture (Table 6.12 and Figure 6.8). The lizards Ctenosaura similis and Iguana iguana are represented by bones from the axial skeleton (vertebrae), forelimbs (ulna and phalanges), hindlimbs (radius, tibia and femur) and pelvis. The lizard´s remains present evidence of burning and irregular fractures (Table 6.12 and Figure 6.9).

Figure 6.8: Element distribution of freshwater turtles expressed in number of identified specimens (NISP)

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Figure 6.9: Element distribution of Iguanas expressed in number of identified specimens (NISP)

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Macrounit 2a

No Bone NISP modification Type Location Burning Action Freshwater turtles (Trachemys & Kinosternon irregular ) Hypoplastron 1 1 fracture missing caudal disarticulation? irregular missing distal articular 1 1 fracture portion disarticulation? irregular missing distal squamosal 1 1 fracture portion disarticulation? irregular missing distal cooking? metacarpal 1 1 fracture portion black smoking? irregular missing distal metapodial 1 1 fracture portion postdepositional peripheal cooking? bone 1 1 scorched smoking? spiral tibia 1 1 fracture shaft defleshing? spiral missing distal ulna 1 1 fracture portion defleshing?

irregular detached vertebrae vertebrae 1 1 fracture epiphysis postdepostional Marine irregular turtles flat bone 3 3 fracture defleshing? (Cheloniidae irregular cooking? ) shaft 3 3 fracture 1 black smoking? 2 missing distal irregular portion, 1 missing cooking? Femur 3 3 fracture proximal portion 1 black smoking? irregular cooking? Lizards metapodial 1 1 fracture shaft black smoking? (Ctenosaura irregular & Iguana) ulna 1 1 fracture shaft disarticulation?

Table 6.12: Reptiles at macrounit 2a: modifications (NISP)

The marine turtles identified here correspond to Chelonia agassiziii (at least 9 individuals) and Lepidochelys (1 individual) (Figure 6.6). Fragmented bones with evidence of spiral fractures, and burning marks are distributed in axial skeleton (vertebrae), forelimbs

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(ulna, radius, metacarpal), hindlimbs ( tibia and metatarsal), skull (squamosal) and the jaws

(dentary and articular) (Figure 6.10).

Lepydochelys usually arrive between Panama and Ecuador to feed from June through

December (Márquez 1990: 45). The Chelonia agassizii inhabits coastal waters of the eastern tropical Pacific Ocean to feed, and is not commonly observed in the open ocean (Márquez

1990: 23).

Figure 6.10: Element distribution of marine turtles expressed in number of identified specimens (NISP)

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Few bones of rodents were found in this unit, and those identified were from

Oryzomys corresponding to a femur fragment. Bats (Chiroptera) are represented by long bones and pelvis with no evidence of modification (Tables 6.13 and 6.14).

Mammalia Specie NISP MNI Biomass cf Carnivora 1 Chiroptera 5 3 Homo sapiens 2 Mammalia 7 Odocoileus 5 3 virginianus Oryzomys 1 1 Procyon cf lotor 2 2 Tamandua cf 1 1 mexicana

Table 6.13: Mammals at macrounit 2a (NISP, MNI and estimated biomass)

Table 6.14: Mammals at macrounit 2a: modifications (NISP).

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An anteater (Tamandua cf Mexicana) was identified in the sample by an ulna fragment presenting cut marks (Figure 6.6). Raccoons were identified by a femur with a v-shaped fracture and dentary from a juvenile individual. At least 2 white-tailed deer were identified corresponding to ribs, tibiae and a juvenile phalange. These bones present spiral fracture and burned marks.

Finally, fragmented human remains from a skull and tibia were also identified. As with the remains of the previous unit, these bones could have been the product of post - depositional disturbance.

Coastal birds are poorly represented in macrounit 2a by 2 ulnae, 1 sternum and a juvenile tibiotarsus (Table 6.15). Two taxa were identified, Meadowlark (Sturnella) and heron (cf Ardea). This last taxon is represented by a bead made from a worked ulna (Table

6.16).

Aves Specie NISP MNI Biomass Aves 1 1 cf Ardea 1 1 1400 cf Aves 1 1 Sturnella 1 1 70 Table 6.15: Birds at macrounit 2a (NISP, MNI and estimated biomass

No Bone NISP modification Type Location Burning Action worked & ulna 1 1 glossy shaft bead missing irregular distal Birds sternum 1 1 fracture portion disarticulation?

Table 6.16: Birds at macrounit 2a: modifications (NISP)

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Macrounit 2b

There are few bones of non-fish vertebrates in this unit. The most common class is again reptiles (Table 6.17). This unit includes lizards such as the black (Ctenosaura similis) and green iguanas (Iguana iguana) and marine turtles, including the green turtle (Chelonia agassizii) and the olive ridley turtle (Lepydochelys cf olivacea).

Reptiles Specie NISP MNI Biomass Ctenosaura similis 13 5 3250 Cheloniidae 1 1 Chelonia agassizii 11 7 245000 Iguana iguana 1 1 Iguanidae 2 cf Cheloniidae 1 cf Testudines 2 Lepydochelys cf olivácea 1 1 50000

Table 6.17: Reptiles at macrounit 2b (NISP, MNI and estimated biomass)

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No Bone NISP modification Type Location Burning Action

skull irregular cooking? fragment 1 1 fracture scorched smoking? spiral femur 1 1 fracture shaft scorched disarticulation?

irregular fracture and missing chopped distal metacarpal 3 3 (heavy blow) portion scorched disarticulation? missing irregular distal Marine ulna 1 1 fracture portion disarticulation? turtles irregular cooking? (Cheloniidae) shaft 1 1 fracture black smoking? irregular missing rib 1 1 fracture distal disarticulation? spiral proximal fracture & portion & femur 1 1 Incision shaft defleshing? irregular Lizards metapodial 1 1 fracture shaft scorched disarticulation? (Ctenosaura irregular cooking? & Iguana) vertebrae 2 2 fracture epiphysis scorched smoking?

Table 6.18: Reptiles at macrounit 2b: modifications (NISP).

Lizards are represented by bones from the axial skeleton (vertebrae, ribs), forelimbs

(ulna, radius), hindlimbs (tibia, and femur), and skull (basisphenoides). In the sample an

Iguana femur has evidence of spiral fracture, incision, and burning (Table 6.18 and Figure

6.12).

Marine turtles are represented by fragments of skull, limb bones (ulna, femur, metacarpal, metatarsal) and ribs with evidence of cut and burning marks (Figure 6.11).

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Figure 6.11: Element distribution of marine turtles expressed in number of identified specimens (NISP) Macrounit 2b.

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Figure 6.12: Element distribution of Iguanas expressed in number of identified specimens (NISP) Macrounit 2b.

Mammals are scarcely present in this unit; there are at least 7 individuals (Figure 6.5).

The most common taxon is bats (Chiroptera) followed by deer (Odocoileus virginianus), opossum (Didelphidae), fox (Urocyon) and Muridae (Table 6.19).

Mammalia Specie NISP MNI Biomass cf Odocoileus 2 2 Chiroptera 4 2 Didelphidae 1 1 Mammalia 3 Muridae 1 1 Odocoileus virginianus 3 1 Urocyon cinereoargenteus 1

Table 6.19: Mammals at macrounit 2b (NISP, MNI and estimated biomass)

There are few bones belonging to bats. Most of the bones are scapula, ulna and humerus and do not presented evidence of being burned or cut. Rodents are scarce too and 166

are represented by a dentary with no evidence of cut or burn marks (Table 6.20). A fox specimen was identified by a premolar. These taxa perhaps are intrusive in this unit.

No Bone NISP modification Type Location Burning Action Medium size mammals (Urocyon, medial Didelphidae irregular portion cooking? & Muridae) dentary 2 2 fracture present 1 black smoking? irregular cooking? Big rib 1 1 fracture shaft 1 scorched smoking? mammals spiral disarticulation (Odocoileus) tibia 3 3 fracture shaft ?

Table 6.20: Mammals at macrounit 2b: modifications (NISP)

At least 3 white-tailed deer individuals, one of them an immature specimen, were identified in this unit. The bone fragments correspond to 2 tibiae, 1 rib, and 1calcaneus.

These bones present spiral fracture and burned marks.

Aves

Specie NISP MNI Biomass

Pelecanus occidentalis 1 1

Table 6.21: Birds at macrounit 2b (NISP, MNI and estimated biomass)

One fragmented radius of a pelican (Pelecanus occidentalis) was found in this unit (Table 6.21).

There is no evidence of cut or burn marks (Figure 6.6).

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Discussion

The non-fish vertebrate remains analyzed at both shelters at Cueva de los Vampiros present evidence of several human activities as well postdepositional processes. The identified taxain this sample represent diverse aquatic and terrestrial ecological conditions around this archaeological site, which also suggests that their exploitation was a local activity. The fauna are characteristic of an estuarine ecosystem and anthropic landscape: muddy zones with a patchy distribution of dry forest, mangroves that house a diversity of species of birds, reptiles and mammals.

With the exception of La Mula points and possibly the unmodified pebbles found at

Cueva de los Vampiros, there is no other artefactual evidence of hunting and processing amphibians, reptiles, birds and mammals. However, an expedient technology surely was used to catch small and big prey. Cooke and Tapia (1994a) mention that nowadays green turtles (Chelonia agassizi) usually are caught by intertidal traps, taking advantage of their feeding behavior.

Based on this small sample, it is premature to draw conclusions about human behavior such as butchery practices, site function, and transport decisions (Faith and

Gordon 2007). Biomass estimates suggest that marine, freshwater turtles, deer and iguanas were an important source of meat. However, lower utility bones such as phalanges, metapodials, mandibles and caparace bones are more common from freshwater and marine turtles than from the other vertebrates such as iguanas and raccoons, suggesting that other

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meaty bones were discarded somewhere else. Cut marks suggest that Vampiros inhabitants were processing turtles and transporting cured meat somewhere else and meanwhile were consuming iguanas and raccoons in the shelter. The burn marks observed on some turtle bones could be related either to the heating process to preserve the meat, or perhaps the bones were thrown on the fire after their meat was eaten off them.

Although fish remains overwhelm other vertebrates, the diversity of the other vertebrates suggests that pre-Columbian inhabitants were engaged in other activities and not only fishing. Reptiles and mammals follow fish in importance. Some of these mammals, bats and rodents are indicative of intrusions. Birds are insignificant in the archaeological record. They are indicative of postdepositional events and/ or their use for ornaments.

Amphibians are also poorly represented in the assemblage and are likely intrusive to these deposits.

Finally, mammals and reptiles (especially marine turtles) contributed the greatest proportion of the biomass to the diet

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Chapter 7

Pre-Columbian Fishing at Cueva de los Vampiros (AG-145)

Introduction

This chapter will describe and analyze the Vampiros fish assemblage. To determine whether the Vampiros deposits resulted from processing fish to retard spoilage and facilitate their transport elsewhere, the analysis focuses on the following: taxonomy (number of fish taxa and their proportion), body part distribution, evidence of physical damage to bones, evidence of differential exposure to heat and cut marks resulting from processing activities.

Recovery Methods and Sampling Bias

As described in Chapter 1, the fish remains for this study come from three 0.5 m2 column samples that test the entire fishing camp deposits. Two were taken from the west wall of

Vampiros-1 (see illustration second chapter). They attained two meters deep, from present ground surface to the base of macrounit 3. One column was taken from the north wall of

Vampiros-2. It reached four and a half meters from modern ground surface. In the field, the remains were collected using 1/8‖ (3.2mm) and 1/16‖ (2mm) meshes. The sediments that passed through the screens were bagged and sieved again at the laboratory through geological screens of 0.25mm, 0.125mm and 0.0625 mm. All fish remains were later analyzed in the lab under a ZEISS stereomicroscope. My original plan/strategy was to analyze all fish

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bones from the three columns. However, while I proceeded to analyze more than 100 samples sieved over 1/8‖ mesh (see Table 7.1 and appendix), I realized that it would be impossible to complete the original work plan. Identification was simply too time-consuming taking into consideration taxonomic diversity. Therefore, I stopped the complete analyses of the columns and decided to examine some samples completely, i.e., identifying and quantifying bones taken over all sieve sizes. Therefore, samples of 1/8‖ from Vampiros-1 and Vampiros-2 were used for general taphonomical aspects (evidence of cut marks, burning) meanwhile completely analyzed samples (samples from 1/8‖ sieve and ¼ of 1/16‖ and 0.25mm) were used to calculate MNI and to estimate relative frequencies of taxa and skeletal completeness. In total, 36 samples were analyzed from macrounit 1, 2a and 2b fromVampiros-1(subunits 5,6,8,13,19,21,27,29 and posthole 5) and 12 samples from

Vampiros-2 (subunits 3, 4, 15, 52, 53, 58 and 59).

Identification and Comparative Collection

Relying on anatomical landmarks, each bone in the Vampiros´ samples was identified to the lowest taxonomic level possible using the comparative fish skeleton collection housed at the

Smithsonian Tropical Research Institute (STRI) in Panama. This collection contains 1570 prepared skeletons belonging to 91families, 221 genera and 356 species from the tropical eastern Pacific and Panamanian freshwater bodies. In this task I was assisted by Máximo

Jimenez, Alexandra Lara and Richard Cooke. Specialized literature was used to avoid confusions on the nomenclature (Betancur and Acero 2004; Betancur., et al. 2007; Cooke and Jiménez 2008; Robertson and Allen 2006; Rojo 1991; Wheeler and Jones 1989).

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Quantification Analysis

In this study, the term "identified" means that fish remains could be attributed to the lowest taxonomic level possible below Superclass, i.e. family, genus or species. Some badly fragmented bones could only be classified as "unidentified", which means that such remains are only recognized as Osteichthyes.

The quantification of data in the present study includes several counting methods that provide information related to element representation, relative frequency of taxa, butchering patterns, dietary contributions and procurement strategies (Grayson 1984; Lyman

2008; Reitz and Wing 2008). Fish data are organized in two groups: 1/8‖ mesh assemblage and fine mesh assemblage. Both groups are summarized using number of identified specimens (NISP), including both complete and fragmented bones. After microscopic observation, each bone was weighed.

The considerable volume of fish bones that occurred at Vampiros rockshelters is typical of Central Pacific sites in Panama because marine shells within these deposits help their preservation. This situation and taxonomic diversity cause considerable sampling problems. In the eastern Tropical Pacific some families have many species, according to

Robertson and Allen (2006); important economic families such as the Carangidae comprise

140 species, the Haemulidae 150, the Sciaenidae 261 and Ariidae 139. Because the standard unit of biological analysis is the species, the identification of species in zooarchaeology is rife with problems. For example, phylogenetically related species often have similar bones; fragmentation affects the precision of identifications. Ontogenetically, the fish skeleton may undergo considerable morphological changes. Although this is a taxonomic ―nightmare‖,

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individual species in these families mentioned above are distributed differentially in near- shore habitats which are informative of the specific habitats that ancient fishermen were exploiting (Bearez 1997; Cooke and Jiménez 2004; Cooke and Jiménez 2008; Cooke 1992a).

Finally, the description of new species in the eastern tropical Pacific makes it necessary to re- evaluate species-level identification. For example a marine catfish found in Parita Bay samples (Notarius cookei) was described in 2002 (Betancur, et al. 2007). By the time I was doing this study new haemulid species were described in Panama (Pomadasys empherus) (Cooke

2009 pers. com.). This is also evident for the use of ―cf‖ on the fish species. Any particular species needs to be seen in context of its comparison to another, but by definition it is not confirmed as the same (Sciades dowi versus Sciades cf dowi)(Reitz and Wing 2008: 36).

The minimum number of individuals (MNI) is calculated for every taxonomic group found in each subunit, taking into account the most abundant anatomical element and size ranges as well (Lyman 2008; Ringrose 1993). The subunits were separations that I defined in the field that I considered analytical units. Therefore, bones from these subunits were considered disconnected from bones of other subunits.

Ideally, the size of individuals should be estimated by allometric predictions which are feasible in Vampiros samples (Reitz and Wing 2008; Wheeler and Jones 1989). Due to time and fund constraints and the taxonomic complexity of these samples, I concentrated on taxonomic and anatomical information demanded by my research questions. I estimated fish size in grams using the proportional method; individual bones were compared with the same element in the STRI reference collection (Cooke, et al. 2004: 215).

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Body part frequency has been calculated to understand the formation processes that created the assemblage, such as fish butchery and processing techniques that may disproportionately deposit cranial and some vertebral elements at processing centers and concentrate other body parts at consumption areas (Belcher 1998; Norton, et al. 1999; Perdikaris 1999;

Perdikaris, et al. 2004; Reitz and Wing 2008; Ringrose 1993).

The relative representation of skeletal elements was calculated according to each anatomic region (olfactory, occipital, investing, etc) for each family (Cannon 1987 and see appendices

1-9). The ratio between cranial (all cranial including atlas –MAU-) and postcranial bones (all vert. & axial plus cleithrum –MAU-) was calculated following Perdikaris et al. (2004). Body part frequency was calculated as follows. First, vertebrate series were determined as the ratio

NISP of each vertebrae type by the number of vertebrae type in a living animal (thoracic, precaudal and caudal) in each family (Figure 7.1 and appendix 1-9).

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Figure 7.1: Fish skeleton nomenclature By Debbi Yee Cannon (1987) and Perdikaris et al (2004)

To evaluate the survivorship of each anatomical region, the survival index was calculated for each bone as a ratio between expected and observed bones. If the survival index (SI) value is bigger than 1, this implies over-representation, while a smaller survival index value (< 1) indicates under-representation. I calculated the expected bone ratio for each family identified and anatomical region. The expected values of each bone were calculated, for each taxa from the ratio of each bone in a complete skeleton multiplied by the total NISP observed in the site (Belcher 1998; Lyman 1994; Perdikaris 1999; Reitz and Wing 2008;

Wheeler 1989; Zohar 2003; Zohar and Cooke 1997; Zohar, et al. 2001a).

The minimal animal units (MAU) have been utilized to understand butchering patterns and post-depositional process in fish assemblages. MAU is calculated dividing NISP of each 175

element by the skeletal part in a living animal (appendix 1-9). The percentage of MAU

(%MAU) is calculated using all MAU values divided by the greatest MAU value in the sample. This percentage should show equal numbers if all elements survive.

Different densities and fragmentation affect the individual survivorship of each bone; therefore, the MAU% of anatomical regions will be use for comparisons rather than individual bones. Several authors have identified numerous problems with NISP, MAU and

MNI (Belcher 1998; Ceron-Carrasco 2005; Grayson 1984; Lyman 2008; Reitz and Wing

2008; Ringrose 1993; Zohar 2003). NISP is affected by differential recovery techniques, butchering practices, aggregation, differential preservation, and sample size among other problems. MNI is influenced by sample size, and difficulties in its calculation, as well as other disadvantages.

Finally, although the skeletal element distribution and MAU are good tools to observe butchering patterns, these are somehow disturbed by the better preservation of robust elements (i.e. dentary, premaxilla, anal pterygipohore, pectoral spines) that give the impression of being the most abundant elements overall.

I acknowledge these difficulties; however, I considered that calculating the MAU values of each family and comparing them with the NISP and MNI would help to assess the importance of certain species and discern butchering patterns from the Vampiros assemblage.

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Taphonomic Analysis

In order to identify butchering practices, taphonomic agents and understand the archaeological processes, several attributes were recorded, and secondary data were calculated as well following Lyman (1994), Belcher (1998) and Zohar (2003; Zohar and

Cooke 1997). The information recorded on the skeletal elements was portion, percentage of completeness and origin of fragmentation. Additionally, the location, number and morphology of the cut marks were also observed.

For Cueva de los Vampiros fish analysis, the samples were evaluated using the fragmentation size classes for each bone as outlined above: these classes were 100%

(complete), <5% (epiphysis), 5- 25 %( less than ¼ complete), 25- 40% (greater than ¼ less than ½ complete), 40-50 %( less than 1/2 less than 3/4 complete), 50-75% (greater than 1/2 than 3/4 less than complete), and 75-90% (broken but still complete) and quantified by number (Figure 7.2).

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Figure 7.2: Fragmentation size classes used for classification of bone state of preservation

Zohar and colleagues (2001a: 1078) standardized the degree of fragmentation using the weighted mean index (WMI) of fragmentation. This index was calculated for each anatomical region to evaluate if the fragmentation patterns observed on fish bones are related to butchering methods and postdepositional damage. Following these authors, I expressed WMI as the mean degree of fragmentation calculated from the relative frequency

(Xi) of each bone in the seven fragmentation size categories and (Wi) the fragmentation category [WMI= (Wi*Xi)/100].

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Encrustation, articulated or fused bones and acid dissolution, as well as cultural processes of heating and trampling of fish bones were registered in the samples (Belcher 1998; Lyman

1994; Zohar 2003; Zohar and Cooke 1997; Zohar, et al. 2001a).

Some studies in salmon (Butler and Chatters 1994) and cod (Nicholson 1992) evaluate the relationship between bone density and their suirvability. Butler and Chatters

(1994) using salmon, mention that scavenging, trampling, sediments chemistry and other processes that occur on human occupation sites would also affect the preservation of proportion of cranial versus postcranial bones. They concluded that that the low density of the ―spongy‖ salmon cranial bones are the reason of the scarcity of cranial bones in archaeological sites in north-western North America .

Nicholson (1992) on the other hand, suggests that density does not have anything to do with the preservation of fish bones. As she points out, if it is removed the human and the animal involvement all fish bones which have suffered similar pre-depositional or postdepositional activities have equal chances to survive. According to her, the shape of the bone is the main factor in determining their survivorship.

Burning on archaeological bones either may represent cooking activities or discarding activities (Nicholson 1993; Roberts, et al. 2002). According to Nicholson (1993), fish bones that achieved different temperatures in an open fire present different colors. A yellowish-red color corresponds to 200ºC, black color on bones indicates temperatures between 300ºC to

500ºC. Bones that present grayish colors were exposed to 600ºC. Finally, whitish bones correspond to temperatures between 700ºC to 900ºC.

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Discussion of Element Distribution, Fractures and Cut Marks from Various Fish Butchering Processes

In this section I will discuss the patterns of element distribution, cut marks and fractures from various butchering processes in order to compare them to Vampiros rockshelters fish analysis.

On medieval contexts in Iceland and Norway, the production of dried fish left either thoracic and pre-caudal vertebrae in the exported fish, or lacked thoracic and most precaudal which would accumulate along with the head and jaw bones at the coastal processing center. The presence of all three vertebral is an indication that at least some whole fish were deposited. If people were consuming fresh fish, evidence from middens would have a balance of skeletal elements more similar to a complete fish skeleton which produce a graph of exactly equal vertebrae proportions for % MAU (33% each)( Krivogorskaya, et al. 2005;

Krivogorskaya., et al. 2005).

The cured fish model assumes that fish were decapitated leaving skull bones and some thoracic, precaudal and anterior caudal vertebrae at a processing site. Meaty elements of the post cranial skeleton such as the supracleithrum, cleithrum, postcleithrum, scapula, coracoid and basipterygium were cured, traded and discarded at the consumer sites. The posterior caudal vertebrae were sometimes left in cured products (Figure 7.3). In this model low frequency of cleithra is indicative of a processing site, whereas their high frequency suggests a distribution of a consumer site (Amundsen 2008; Barrett 1997; Hoffman, et al.

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2000; Høtje 2006; Krivogorskaya, et al. 2005; Krivogorskaya., et al. 2005; Norton, et al. 1999;

Perdikaris 1999; Stewart and Gifford-Gonzalez 1994).

Figure 7.3: The cured fish model (photo Y. Krivogorskaya)

Aulin Tourunen (2008) analyzed a fish bone assemblege from a medieval context at

John Street, Waterford, Ireland. Based on NISP, MAU counts and element distribution and graphic distribution, Tourunen suggests that the preponderance of the hyoid arch, branchial apparatus and pectoral girdle which she called the ―throat‖ in comparison to other regions were result of removing and discarding activities when gutting the fish. She did not mention fractures or cut marks.

Other studies considered butchering methods in preserving fish are based on size.

In a processing site small fish are butchered keeping the skull intact and using a ventral

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midline cut; gills (operculum: opercular, preopercular, subopercular, interopercle and branchial skeleton: ceratohyal, epihyal, hypohyal, urohyal, branchiostegal rays) and guts are removed and discarded. Large fish are split dorsally along the vertebral column and skull; fragmented cranial bones also are discarded (Belcher 1998; Zohar 2003; Zohar and Cooke

1997; Zohar, et al. 2001). These ideal models could be affected by cultural practices or differential preservation of skeletal elements as I mentioned above (Barrett 1997; Nicholson

1996; Nicholson 1993; Zohar, et al. 2001a).

Belcher (1998) in his ethnoacraheological study at Baluchistan and Indus Valley mentions that fish is butchered depends of their size and morphology. If people have a direct access to fresh fish at the household village, they eviscerate and remove the gill arch region for both small and large fish. The cut marks are distributed on the lateral side of the bones on the gill arch region. The bone assemblage of fresh fish processing areas is characterized for bones from the fins, gills and tails (Belcher 1998: 176). If people have an indirect acces to fish at the household village, gutted small fish are transported whole and cooked. Cranial remains from clupeids and mugilids are consumed. The large size fish like catfish are butchered by gutting and removing the gills. Heads are chopped in half along the medial line. Pectoral and dorsal spines are broken near the proximal articular head to avoid injuries, therefore on the archaeological site proximal or medial fragments are found. Cut marks present on large fish as chop-marks and are distributed on cranial elements (Belcher 1998: 170). Cut marks laterally located on cranial bones and first vertebrae set are associated with removal of the head. No cut marks are observed on small fish (Belcher 1998:172).

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In a commercial context, for example, to trade from coastal to inland villages, fish are gutted and the entrails are discarded. For large fish, the body and head are split laterally from the ventral surface along the midline for ease in drying and saltying. The neurocranium is cut in half; incisions are placed on postero-medial region of the flanks to introduce salt and the posterior-medial region of the lateral facial area present cut marks to allow the head lying flat. Smaller fish is dried whole. Cut marks occur along the medial surface of cranial elements. The branchiostegal region and otoliths added into the archaeological record

(Belcher 1998: 175). In Belcher ´s (1998) ethoarchaeological study some missing areas such as the branchial arch and caudal vertebrae suggests indirect obtention and were disposed in a processing area.

Willis et al. (2008) mention that cut marks on fish are rarely reported because their of their low visibility. They butchered 37 fish using stone tools and a metal knife following methods provided in ethnographic accounts and by modern fish processors. The authors butchered

37 fish.

The first method:

―..a fish is placed on its side and an incision is made from the anus to the pectoral

fins. Next, the head and viscera are removed. The initial ventral incision is extended

from the anus to the caudal fin and the fish is laid open and flat with the vertebral

column exposed (Figure 7.4). Cuts are then made laterally on both sides of the

vertebral column, severing rib attachments. Finally, the vertebral column and caudal

fin are cut off of the remaining fillets (Willis et al 2008:1439)”.

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The second method:

―a local fish market employee,was used to butcher the morphologically distinct

summer flounder. After placing the flounder on its blind side, the initial incision is

made along the length of the dorsal fin, from the pectoral girdle to the caudal fin.

Following the initial incision, long strokes are used to separate away the fillets. The

same incision is made along the anal fin, and long strokes are used to remove the

ventral portion of the fillet. A cut is made posterior to the pectoral girdle to

completely remove the fillet. The head and viscera are removed. Finally, placing the

flounder on its ocular side, the same process is used to remove the blind-side fillet.

The vertebral column, ribs, and dorsal, anal, and caudal fins remain articulated (Willis

et al 2008:1439-40)‖.

The cut marks produced by filleting fish using these two methods are present not only on diagnostic but on undiagnostic bones; these were mostly the vertebral neural and haemal spines, transverse processes, ribs and pterygiophores these were mostly the vertebral neural and haemal spines, transverse processes, ribs and pterygiophores. According to Willis et al

(2008:1441) cut marks on the vertebral complex are a butchering mistake made when people decapitated catfish.

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Figure 7.4: Fish filleting (Peebles et al 2010)

Similar to Belcher´s work, Zohar and collegues (1997, 2001; 2003) found that in contemporary communities in Panama, butchering practices differ according to the size of the fish. She describes two methods of processing the fish. The first method is used in fish larger than 35 cm (<400gr) which were butchered dorsally through their skull (Figure

7.5). In the second method, used in fish smaller than 35 cm (>400gr), the specimens were processed whole and gutted ventrally. In large fish as illustrated in Table 7.1, several cranial bones will be lost. In addition, cranial bones that are situated along the longitudinal cut are frequently damaged and therefore have slight opportunities to survive.

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Figure 7.5: Fish butchered by method 2 (left- center) and method 1(right) (Zohar and Cooke

1997).

Method 1 /Small fish (<400gr) Method 2/Large fish(>400 gr) The specimens were processed whole and Butchered dorsally through their skull. gutted ventrally Damage: Neurocranial bones intact. Damage: Fractures are found at the Damage is found at the coracoid, cleithrum cleithrum, coracoid, premaxilla, quadrate, and basypterigium. metapterygoid, epihyal, preopercle, interopercle and urohyal . Missing: None Missing: First dorsal spine and supraoccipital´s crest in Ariidae are missing.

Table 7.1: Butchering methods describe by Zohar (2003), Zohar and Cooke (1997) and Zohar et al (2001)

Butchery analysis on fish bones from an archaeological perspective has depended on categorizations (Belcher 1998; Zohar 2003) . This is complemented by studies that aim to

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identify what the marks on the surface of the bones represent (Belcher 1998; Fisher 1995;

Seetah 2006; Willis et al 2008).

A bone with recent damage has sharp edges, and a clean, white appearance or light color that contrasts with the darker color of the bone's cortical surface. Bones with ancient damage lack of these characteristics (Table 7.2).

Damage

Predepositional Postdepositional break with angular break with smooth borders, borders and light same color overall color

Table 7.2: Distintion between predepositional and postdepositional

As Willis et al. (2008) mention , cut marks on fish are rarely reported because their of their low visibility. They are present not only on diagnostic but on undiagnostic bones. Following

Belcher´s (1998), Fisher (1995), Seetah (2006) and Zohar´s (2003), I observed each bone to identify cut marks, recording its location, number and morphology (Table 7.3).

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Cut Marks and Fractures Characteristics Position Morphology Direction Postdepositional light color lateral irregular Longitudinal

Predepositional same color overall Mid impact Transversal crushed rough entry and fracture exit points Medial line chopped smooth entry and fractured exit points Posterior/Caudal "straight" cut fine transversal and shallow incision Anterior/Cranial impact circular perforation Dorsal/Superior Ventral/Inferior

Table 7.3: Criteria for identying cutmarks and fractures

The majority of cut marks tended to be shallow and small striations (cut) caused by knife.

Impact is like a percussion pit with a circular perforation or depresion for using a hammerstone or perhaps a polished axe. Crushing is a splintererd marging of the bone from a hammerstone or perhaps a polished axe, and chopping mark is smooth entry and fractured exit points for using a hammerstone or perhaps a polished axe.

Fish Families and Environment

Of the 89957 fish bones recovered from 1/8‖ and fine meshes (1/16‖ and 0.25mm), 14490 were identified to at least family level. Significantly, more bones were recovered in

Vampiros-1 than Vampiros-2 (Table 7.4). The proportion of identified specimens in each shelter was different (see Table 7.5). At Vampiros-2 poor preservation did not allow retention of enough morphological features and some fragments were allocated to a family level. Both shelters account for 40 families, 91 genera and at least 135 species.

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Locus Identified Unidentified Total Burnt Total NISP NISP % NISP % Vampiros-1 10987 65.2 5865 34.8 5275 16852 (1/8”mesh) Vampiros-1 2610 7.5 92.5 21238 34965 (fine mesh) 32355 Vampiros-2 420 36.2 740 63.8 888 1160 (1/8”mesh) Vampiros-2 473 1.3 36507 98.7 35020 36980 (fine mesh) Total 14490 75467 62421 89957

Table 7.4: (NISP) and percentage of fish remains identified and unidentified by loci

Using the measure of NISP, the relatively most abundant family from the 1/8‖ mesh of the complete analyzed samples is Carangidae (25.1%) followed in turn by Ariidae (15.3%),

Tetraodontidae (7.9%), Sciaenidae (5.4%), Polynemidae (5.4%), Haemulidae (5%),

Pristigasteridae (3.4%), Belonidae (2.4%), Clupeidae (2.1%), Ephippidae (1.2%) and

Albulidae (1.1%) . Other families are represented by less than 1% (see Table 7.2).

The situation for the sample recovered from 1/16‖ mesh is slightly different. Here the most abundant is Carangidae (16.2%) followed in turn by Ariidae (13.4%), Sciaenidae (4.9%),

Tetraodontidae (4.6%), Clupeidae (4.6%), Polynemidae (3.4%), Belonidae (3.1%),

Haemulidae (2.8%), Pristigasteridae (2%), Mugilidae (0.9%) and Centropomidae (0.8%). The remaining families are represented by less than 1% (see Table 7.3).

Among these families, small specimens from Engraulidae, Hemiramphidae and

Stromatidae are more frequent from 1/16‖ mesh than 1/8‖ mesh. A similar distribution is observed on the additional material recovered on 1/8‖ mesh (Table 7.6).

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Macrounit Family 1 2a 2b Total 1/8" Total 1/16" Mesh Class 1/8" 1/16" 0.25 1/18" 1/16" 0.25 1/18" 1/16" 0.25 % Rank % Rank Shelter V1 V2 V1 V2 V1 V2 V1 V2 V1 V2 V1 V2 V1 V2 V1 V2 V1 V2 Achiridae B 1 7 0.1% 0 0.0% Albulidae B 6 1 1 44 14 5 3 56 1.1% 10 19 0.6% Ariidae B 159 1 62 8 582 227 40 251 2 60 42 750 15.3% 2 439 13.4% 2 Batrachoididae B 2 1 18 2 2 26 0.5% 3 0.1% Belonidae B 21 7 3 89 62 2 39 23 5 116 2.4% 7 102 3.1% 6 Carangidae B 462 135 3 759 238 35 302 4 107 13 1231 25.1% 1 531 16.2% 1 Carcharhinidae C 2 3 5 1 11 0.2% 1 0.0% Centropomidae B 9 1 3 1 18 15 1 14 1 4 34 0.7% 25 0.8% 10 Clupeidae B 36 29 1 63 48 15 21 49 9 105 2.1% 8 151 4.6% 4 Cynoglossidae B 1 6 0.1% 1 0.0% Dasiatidae C 1 1 6 0.1% 2 0.1% Eleotridae B 1 2 2 9 0.2% 2 0.1% Elopidae B 1 6 0.1% 1 0.0% Engraulidae B 2 1 7 12 6 3 15 0.3% 22 0.7% Ephippidae B 10 43 8 26 10 59 1.2% 9 18 0.5% Erythrinidae B 1 6 0.1% 1 0.0% Fistulariidae B 2 6 3 2 8 0.2% 8 0.2% Gerridae B 1 1 9 3 1 2 3 1 16 0.3% 9 0.3% Gobiidae B 1 6 0.1% 1 0.0% Ginglymostomatidae C 1 7 0.1% 0 0.0% Haemulidae B 92 28 1 148 47 6 53 9 1 246 5.0% 5 92 2.8% 7 Hemiramphidae B 5 1 2 2 3 11 0.2% 6 0.2% Lobotidae B 5 3 1 11 0.2% 3 0.1% Lutjanidae B 1 4 1 11 0.2% 1 0.0% Mugilidae B 6 1 6 3 34 12 1 30 4 2 47 1.0% 28 0.9% 9 Muraenidae B 3 1 6 0.1% 4 0.1% Muraenesocidae B 1 7 0.1% 0 0.0% Narcinidae C 1 6 0.1% 1 0.0% Ophichthidae B 1 7 0.1% 0 0.0% Paralichthyidae B 2 4 5 2 1 2 12 0.2% 10 0.3% Polynemidae B 77 14 207 39 19 84 36 5 290 5.9% 4 113 3.4% 5 Pristigasteridae B 49 20 110 32 6 30 8 1 165 3.4% 6 67 2.0% 8 Sciaenidae B 75 28 7 206 77 8 104 36 4 287 5.9% 4 160 4.9% 3 Scombridae B 17 1 5 7 1 24 4 28 0.6% 13 0.4% Serranidae B 1 2 1 8 0.2% 2 0.1% Sphyraenidae B 17 29 3 1 4 1 52 1.1% 10 5 0.2% Sphyrnidae C 1 1 2 4 7 0.1% 7 0.2% Sternopygidae F 1 7 0.1% 0 0.0% Stromateidae B 1 2 1 7 0.1% 3 0.1% Tetraodontidae B 70 12 308 120 5 128 1 14 385 7.9% 3 151 4.6% 4 Urotrygonidae C 1 6 0.1% 1 0.0% Osteichthyes 264 275 11 538 657 147 183 6 152 40 31 814 16.6% 1282 39.0% Grand Total 1384 4 627 45 0 0 3247 0 1648 299 0 0 1313 13 537 129 31 0 4900 100.0% 3285 100.0% Table 7.5: Complete analyzed units: (NISP) and percentage of identified fish remains by mesh, shelter and macrounit including the rank (in red) most abundant families. C (Cartilaginous fish- Chondrichthyes), B (Bony fish –Osteichthyes).

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Macrounit Family 1 2a 2b Grand Total % Mesh Class 1/8" 1/18" 1/18" Shelter V1 V2 V1 V2 V1 V2 Albulidae B 1 9 3 34 47 0.9% Ariidae B 10 1 44 1 78 134 2.6% Batrachoididae B 1 13 14 0.3% Belonidae B 3 20 50 73 1.4% Carangidae B 182 5 242 121 1253 35 1838 35.3% Carcharhinidae C 1 2 3 0.1% Centropomidae B 1 1 2 4 0.1% Clupeidae B 29 1 12 140 182 3.5% Engraulidae B 2 2 0.0% Ephippidae B 6 22 9 131 168 3.2% Fistulariidae B 2 2 0.0% Gerridae B 16 2 18 0.3% Haemulidae B 1 31 11 43 0.8% Lobotidae B 2 2 0.0% Mugilidae B 5 51 19 119 2 196 3.8% Ophichthidae B 1 1 0.0% Paralichthyidae B 1 1 0.0% Polynemidae B 33 2 108 73 437 15 668 12.8% Pristigasteridae B 15 57 5 119 196 3.8% Sciaenidae B 4 4 11 17 36 0.7% Scombridae B 8 11 18 95 132 2.5% Serranidae B 7 7 0.1% Sphyraenidae B 4 14 16 26 3 63 1.2% Tetraodontidae B 82 4 169 45 790 15 1105 21.2% Osteichthyes 11 2 169 97 279 5.4% Grand Total 396 980 3436 5214 100.00%

Table 7.6: Supplementary analyzed units: (NISP) and percentage of identified fish remains by mesh, shelter and macrounit.

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Figure 7.6: Frequent genera 1/8‖ mesh (excluded genera < 1%)

Figure 7.7: Frequent genera 1/16‖ mesh (excluded genera <1%)

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Among these families, the commonest genera found in both shelters are Selene sp (21.5%),

Polydactylus sp (7.9%), Guentheridia (7.8%), Ariopsis (6.1%), Cathorops (5.8%), Cynoscion sp

(4.3%), Ilisha (4.1%), Caranx sp(4.0%), Notarius sp (3.7%), Orthopristis sp (3.3%), and

Opisthonema sp(2.6%) on 1/8‖ mesh (Figure 7.6).

In the case of 1/16‖ (Figure 7.7), the most frequent genera are Selene sp (22.2%), Opisthonema sp (9%), Sphoeroides sp (6.9%), Polydactylus sp(6.8%), Ariopsis (5.9%), Cathorops sp (4.6%), Ilisha

(3.6%), Tylosurus sp(3.5%), Orthopristis (3.2%) and Chloroscombrus (2.6%).

Graphs 7.8 and 7.9 show the relative abundance of the species in 1/8‖ and 1/16‖ meshes. So far, Pacific moonfish (Selene peruviana) is the commonest species found in both shelters. This speciesis followed by spotted puffer (Guentheridia formosa), tete sea-catfish

(Ariopsis seemani ), yellow bobo (Polydactylus opercularis), hachet herring (Ilisha furthii),

(Orthopristis chalceus), Pacificthread herring (Opisthonema libertate), racer croaker (Ophioscion typicus), Pacificcrevalle-jack (Caranx caninus), Pacific bumper (Chloroscomus orqueta) and

Kesslers sea catfish (Notarius kessleri).

The presence of these species gives an idea of the environment surrounding Cueva de los Vampiros at the time of its occupation. The Pacificmoonfish (Selene peruviana) avoids turbid estuarine waters but usually lives in schools in clear water on rocky and sandy substrates. Additionally, the following species such Pacific sierra (Scomberomorus sierra), green jack (Caranx caballus,) barracuda (Sphyraena ensis), bonefish (Albula spp) and needlefish

(Tylosurus cocodrilus fodiator) are also not common in turbid estuarine waters. These species are available in the clear water column not far from the coast. These clear-water species are more frequent in Macrounit 2b than in Macrounit 2a and 1. The marine environmental

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conditions surrounding the Santa Maria River´s delta and Vampiros shelters appear to have been different from the present-day conditions since the habitat described corresponds to clear waters found at the outer margin of the mixing plume. Another possible scenario may be that Pre-Columbian inhabitants obtained fish at some distance from Vampiros (Carvajal

Contreras, et al. 2008).

On the other hand, some species that prefer turbid estuarine conditions such as tete sea-catfish (Ariopsis seemanni), congo sea-catfish (Cathorops spp), Kesslers´ sea- catfish (Notarius kessleri), hachet herring (Ilisha furthii), racer croaker (Ophioscion typicus) and whitefin weakfish

(Cynoscion albus) increase from the lower macrounit to the upper macrounit.

This temporal variation of fish species is correlated to changes previously documented in the coastal geomorphology that occurred since the Holocene. The Santa Maria River´s oscillations may well have been a factor (Barber 1981; Clary, et al. 1984; Dere 1981).

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Figure 7.8: The relative abundance of the species in 1/8‖ mesh at Vampiros 1 and 2. 195

Figure7.9: The relative abundance of the species in 1/16‖ mesh at Vampiros 1 and 2. 196

Figure 7.10: The most frequent species from 1/8‖ mesh(%NISP excluding < 1%)

In terms of NISP (see Tables 7.7 to 7.9 and Figures 7.10 and 7.11), the most abundant species at Vampiros is the Pacificmoonfish (Selene Peruviana). Its distribution in

Vampiros is similar to other Pre-Columbian sites such as Cerro Juan Diaz (Jiménez 1999;

Jiménez and Cooke 2001) where Pacificmoonfish was also the most abundant species. This is the third species in order of importance at Sitio Sierra (Cooke and Ranere 1999). The second most abundant species at both shelters is the puffer fish (Guentheridia formosa), which is very abundant at Vampiros in comparison to other sites on Parita Bay (Cooke and Jiménez

2004; Cooke 1992a; Isaza-Aizpurua 2007). The bullseye puffer (Sphoeroides annulatus) is the eight species in order of importance. Both puffer fish (Guentheridia formosa and Sphoeroides annulatus) have toxic guts and are not eaten in Panama in modern times. However, their abundance especially in Vampiros and other archaeological sites such as La Mula –Sarigua

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and Cerro Juan Diaz along Parita Bay could suggest that they were used for food (Cooke and

Jiménez 2004; Cooke 1992a; Cooke and Tapia-Rodríguez 1994a; Isaza-Aizpurua 2007).

Today bullseye (Sphoeroides annulatus) is consumed in Mexico and has been evaluated for aquaculture (García-Ortega, et al. 2002; Guzmán 2008; Nunez-Vazquez, et al. 2000; Stahl

2003: 186). Gilbert and Starks (1904) mention that bullseye puffer (Sphoeroides annulatus) and spotted puffer (Guentheridia formosa) were common and were abundant in Panamanian markets. Today, Panamanians consider puffer fish disgusting (Martin-Rincon, personal communication 2007). There are other perceptions of this fish family worldwide. According to Titcomb (1972), people in Hawaii did not eat puffer fish frequently. However, he says that if the gall-bladder and the skin are removed carefully, the fish can be eaten if it is previously cooked. He also includes a note about how to identify a poisonous Tetraodontid fish: ―the malani is a poisonous fish indeed. The ‗o‘opuhue manalao, however, is edible to tell them apart look at the teeth. If the teeth are yellow the fish is poisonous. But if the teeth are bright, clear white the ‗o‘opuhue is good to eat‖(Titcomb 1972:131).

Following in order of importance at Vampiros are the species tete sea-catfish

(Ariopsis seemani), yellow bobo (Polydactylus opercularis), hachet herring (Ilisha furthi) , humpback grunt (Orthorpristis chalceus), Pacific bumper (Chloroscombrus orqueta), green jack ( Caranx caballus), blue bobo (Polydactylus approximans), Steindachner's sea catfish (Cathorops steindachneri ), kesslers´ sea-catfish (Notarius kessleri), Pacificthread herring (Ophistonema libertate), white mullet (Mugil curema), crocodile needlefish (Tylosurus c. fodiator), Pacificcrevalle- jack ( Caranx caninus), racer croaker (Ophioscion typicus) and Shortjaw leatherjack (Oligoplites refulgens) (Figures 7.10 and 7.11) .

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Figure 7.11: The most frequent species from 1/16‖ mesh (%NISP excluding < 1%)

Surprisingly few sharks were observed in the sample greater than 1/8‖ mesh. The following species were found from 1/16‖ mesh: chondrytes like the scalloped hammer head shark (Sphyrna lewini), giant electric ray (Narcine entemedor) and stingray (Urotrygon asterias).

Bony fish species were identified, including pike needle fish (Strongylura exilis), and smaller animals such as big-scale anchovy (Anchovia macrolepidota), bigeye corvine (Isopisthus remifer),

Pacificdrum (Larimus pacificus), vermiculate croaker (Ophioscion cf vermicularis) and angel croaker

(Paralonchurus cf goodei); these species are not common in the shallow waters of Parita Bay

(Figures 7.8 and 7.9).

A few freshwater species were identified at Vampiros rockshelters: a single caudal vertebra of knifefish (Sternopygus sp) and a vertebra of tiger fish (Hoplias microlepsis). These could be the result of an accidental catch by the Pre-Columbian inhabitants. Another

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possibility is that the specimens were carried from the Santa Maria River´s current to the coast. In other words, their low representation on archaeological samples is the result of natural conditions, suggesting that freshwater fish were not an important resource in

Vampiros.

The presence of Pacific thread herring, especially in 1/16‖ mesh, (Opisthonema liberate) suggests that people were fishing near sandy beaches using fine mesh nets. As observed in the graphs (Figures 7.8 to 7.11), this group is not abundant, which means that the natural conditions surrounding Vampiros were not dominated by sandy beaches. Anchovies such as big-scale anchovy (Anchovia macrolepidota) and Pacific sabre-tooth anchovy (Lycengraulis poeyi ) are scarce in Vampiros samples. This situation contrasts with the abundance of these species nowadays, which live close to the coast, particularly at the end of dry season (Cooke and

Tapia-Rodríguez 1994a). These species were identified at Cerro Juan Diaz (Jiménez 1999) during Pre-Columbian times. Similar behaviour during the dry season is observed also for

Pacific spadefish (Chaetodipterus zonatus) and Panama spadefish (Parapsettus panamensis), which are more abundant in Vampiros than other sites on Parita Bay (Cooke 2001; Jiménez and

Cooke 2001).

The following Table 7.5 summarizes the habitat composition of the ten most abundant species and the probable method of fishing based on biological and archaeological literature (Cooke and Ranere 1999; Cooke 1988, 1992a, 2004b; Cooke and Tapia-Rodríguez

1994a; Jiménez and Cooke 2001). All the fishes are estuarine or enter occasionally the estuaries or lagoons. There is a poor representation of freshwater and sandy beach species.

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Ecology Specie Probable fishing Method Regular at middle estuary + mangrove Guentheridia formosa Clossing off esterillo outlets Regular at middle estuary + mangrove Selene peruviana Fine meshed gill nets Regular at middle estuary + mangrove Sphoeroides annulatus Clossing off esterillo outlets Regular at middle estuary + mangrove Polydactylus opercularis Tidal weirs Regular at middle estuary + mangrove Ariopsis seemani Clossing off esterillo outlets Fine meshed gill nets/ clossing off Regular at middle estuary + mangrove Chloroscombrus orqueta esterillo outlets Regular at middle estuary + mangrove Ilisha furthi Clossing off esterillo outlets Ocassional at middle estuary + mangrove and bar-formed Lagoons Caranx caballus Probably clossing off esterillo outlets Regular at middle estuary + mangrove Opisthonema libertate Fine meshed gill nets Clossing off esterillo outlets Mugil curema Enter gill nets by encircling Regular at middle estuary + mangrove Orthoprisitis chalceus Tidal weirs Ocassional at middle estuary + mangrove and bar-formed Lagoons Caranx caninus Probably clossing off esterillo outlets Ocassional at middle estuary + mangrove and bar-formed Lagoons Sphyraena ensis Probably clossing off esterillo outlets Regular at middle estuary + mangrove Notarius kessleri Clossing off esterillo outlets Regular at middle estuary + mangrove Ophiscion typicus Clossing off esterillo outlets Shallow waters off the coast Chaetodipterus zonatus More active during the summer Table 7.7: The ecological composition of the most abundant species and the probable method of fishing (based on Cooke 1992).

As mentioned before, there are no archaeological materials allowing the inference that people at Vampiros were using sink nets. However, it is probable that they were using intertidal traps and fine-meshed gill nets using perishable materials. The available information above suggests that people were fishing close to the coast in habitats with a weak freshwater influence such as the middle estuary and mangrove with occasional access to more clear water conditions, perhaps during the dry season.

Diversity and Equitability

To examine similarities and/or differences in the fish assemblage at Vampiros, I compared its three stratigraphic units following some ecological quantitative methods used by zooarchaeologists (Bearez 1996; Lyman 2008; Reitz and Masucci 2004; Reitz and Wing

2008). Diversity and equitability as a I mentioned for the mollusc chapter, allows to

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evaluate subsistence strategies in terms of the variety of fish used by pre-Columbian inhabitants and the ―the evenness with which these resources are used‖ (Reitz and Massuci

2004:79). Differences in these two indexes are result of natural environment, human agency, season of site occupation, and social positioning (Reitz & Wing 2001). Both indexes are calculated for MNI and Biomass. The information was obtained from Tables 7.8 to 7.10.

Diversity is calculated using Shannon Index as follows:

ni The number of individuals in a species i; the abundance of species i.

S The number of species.

N The total number of all individuals

pi The relative abundance of each species, calculated as the proportion of individuals of a given species to the total number of individuals in the community:

. Low=0 -1.5 Moderate= 1.5-3.5 High=3.5- 5.0

Equitability is obtained using Shannon function as S is the natural log of the number

of species for which MNI was estimated, and H‘ is the Shannon-Weaver function

(Lyman 2008;. Reitz and Masucci 2004: 79).

V´= H´/loge S

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Low=0 -0.25 Moderate= 0.25-0.75 High=0.75- 1

Macrostrata MNI #Taxa Diversity Equitability M1 375 123 1.531 0.258 M2a 901 178 1.128 0.166 M2b 539 127 0.487 0.077 Table 7.8: Diversity an equitability at Vampiros 1 and 2 by MNI

Macrostrata Biomass Kg #Taxa Diversity Equitability M1 108.365 123 1.727 0.369 M2a 297.66 178 1.338 0.235 M2b 181.935 127 0.677 0.130 Table 7.9: Diversity an equitability at Vampiros 1 and 2 by biomass

In terms of MNI and biomass (Tables 7.8 and 7.9), diversity and equitability are low at the beginning of the occupation (macrounit 2b) and slightly higher towards the end of the occupation (Macrounit 1). In general both indexes indicate a wide range of fish taxa are present in Vampiros that contribute to provide meat.

Dominance and Contribution to Diet

Dominance can be observed using the percentage of MNI for each taxon and its contribution to the Pre-Columbian diet. In the following Tables (7.10 to 7.12), the most frequent taxa in each macrounit are summarized.

As it was mentioned above, MNI is calculated for every taxon taking into account the most abundant anatomical element and size ranges. On the other hand, the Estimated

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Biomass is the sum of the inferred body masses of all individuals of each taxon (Cooke and

Ranere 1999). In the following Tables, I will consider each macrounit separately. I will provide NISP, NISP%, MNI, MNI% and Estimated Biomass.

In Macrounit 1 (Table 7.10), Pacificmoonfish (Selene peruviana) represents 20.04 %

MNI (in red bold), followed by tete sea-catfish (Ariopsis seemani) (4 %), spotted puffer

(Guentheridia formosa) (4.27%), bullseye puffer (Sphoeroides annulatus) (2.93%), brassy grunt

(Orthopristris chalceus) (4.55%) and Pacific bumper (Chloroscombus orqueta) (4.27%). In contrast, the percentage of Estimated Biomass (in red bold also) indicates that the Kesslers´ catfish

(Notarius kessleri) contributed a larger proportion of the diet (5.43%), followed by Red sea- catfish (Bagre pinnimaculatus) (5.12%), spotted puffer (Guentheridia formosa) (4.96%),

Pacificmoonfish (Selene peruviana) (4.74%), and yellow bobo (Polydactylus opercularis) (4.42%).

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Family genus Species NISP % MNI % Biomass (gr) % Range Albulidae Albula esuncula 8 0.53% 2 0.53% 800 0.74% 400 Ariidae Ariopsis seemani 74 4.91% 15 4.00% 3685 3.40% 5-650 Ariidae Bagre 1 0.07% 1 0.27% 250 0.23% Ariidae Bagre panamensis 11 0.73% 3 0.80% 900 0.83% 150-1450 Ariidae Bagre pinnimaculatus 4 0.27% 4 1.07% 5550 5.12% 300-2500 Ariidae Cathorops cf fuerthii 2 0.13% 2 0.53% 190 0.18% 50-140 Ariidae Cathorops fuerthii 20 1.33% 11 2.93% 2470 2.28% 40-450 Ariidae Cathorops multiradiatus 1 0.07% 0.00% 0.00% 180 Ariidae Cathorops 21 1.39% 7 1.87% 1300 1.20% 70-270 Ariidae Cathorops cf steindachneri 3 0.20% 2 0.53% 360 0.33% 140-220 Ariidae Cathorops steindachneri 13 0.86% 8 2.13% 1280 1.18% 50-300 Ariidae Cathorops tuyra 1 0.07% 1 0.27% 80 0.07% Ariidae 50 3.32% 0.00% 0.00% Ariidae Notarius cf cookei 1 0.07% 1 0.27% 3500 3.23% Ariidae Notarius kessleri 14 0.93% 10 2.67% 5880 5.43% 100-2000 Ariidae Notarius 8 0.53% 2 0.53% 1550 1.43% 150-1400 Ariidae Notarius planiceps 2 0.13% 1 0.27% 180 0.17% Ariidae Notarius troschellii 3 0.20% 3 0.80% 3900 3.60% 800-1800 Ariidae Sciades dow i 1 0.07% 1 0.27% 2000 1.85% Batrachoididae Batrachoides boulengieri 3 0.20% 2 0.53% 380 0.35% 130-250 Belonidae Strongylura cf exilis 5 0.33% 1 0.27% 80 0.07% 80-600 Belonidae Strongylura exilis 1 0.07% 0.00% 0.00% 350 Belonidae Strongylura 1 0.07% 0.00% 0.00% 180 Belonidae Strongylura cf scapularis 1 0.07% 0.00% 0.00% 350 Belonidae Strongylura scapularis 4 0.27% 0.00% 0.00% 100-140 Belonidae Tylosurus cocodrilus fodiator 18 1.19% 4 1.07% 1750 1.61% 120-800 Carangidae Caranx cf caballus 1 0.07% 1 0.27% 300 0.28% Carangidae Caranx caballus 13 0.86% 5 1.33% 1120 1.03% 100-400 Carangidae Caranx caninus 8 0.53% 6 1.60% 2450 2.26% 100-850 Carangidae Chloroscombrus orqueta 56 3.72% 16 4.27% 1220 1.13% 15-150 Carangidae Hemicaranx leucurus 6 0.40% 2 0.53% 270 0.25% 100-200 Carangidae Hemicaranx 1 0.07% 0.00% 0.00%

Table 7.10: Vampiros 1 and 2 : Fish species list at Macrounit 1

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Family genus Species NISP % MNI % Biomass (gr) % Range Carangidae Hemicaranx zelotes 1 0.07% 1 0.27% 200 0.18% Carangidae 47 3.12% 0.00% 0.00% Carangidae Oligoplites altus 1 0.07% 0.00% 0.00% 60 Carangidae Oligoplites cf altus 1 0.07% 0.00% 0.00% 190 Carangidae Oligoplites 8 0.53% 2 0.53% 0.00% 100-130 Carangidae Oligoplites refulgens 37 2.46% 9 2.40% 1165 1.08% 60-250 Carangidae Oligoplites saurus inornatus 1 0.07% 1 0.27% 190 0.18% Carangidae Selene 117 7.76% 3 0.80% 0.00% Carangidae Selene peruviana 302 20.04% 50 13.33% 5140 4.74% 20-250 Carcharhinidae Carcharhinus leucas 1 0.07% 0.00% 0.00% 2000 Carcharhinidae Negaprion brevirostris 1 0.07% 0.00% 0.00% 3000 Centropomidae Centropomus cf armatus 1 0.07% 1 0.27% 120 0.11% Centropomidae Centropomus armatus 2 0.13% 2 0.53% 270 0.25% 120-150 Centropomidae 1 0.07% 0.00% 0.00% Centropomidae 2 0.13% 0.00% 0.00% Centropomidae Centropomus 6 0.40% 3 0.80% 630 0.58% 110-2500 Centropomidae Centropomus unionensis 2 0.13% 1 0.27% 40 0.04% 40-120 Clupeidae Opisthonema cf libertate 25 1.66% 2 0.53% 300 0.28% 80-200 Clupeidae Opisthonema libertate 41 2.72% 8 2.13% 1050 0.97% 80-250 Dasiatidae Dasiatys longa 1 0.07% 0.00% 0.00% 3000 Eleotridae Eleotris picta 1 0.07% 0.00% 0.00% 75 Engraulidae Anchoa 1 0.07% 0.00% 0.00% 15 Engraulidae Anchovia macrolepidota 2 0.13% 1 0.27% 40 0.04% Ephippidae 1 0.07% 0.00% 0.00% Ephippidae Chaetodipterus zonatus 3 0.20% 1 0.27% 500 0.46% Ephippidae cf Chaetodipterus zonatus 1 0.07% 1 0.27% 700 0.65% Ephippidae Parapsettus panamensis 5 0.33% 2 0.53% 380 0.35% 130-300 Gerridae Eucinostomus currani 1 0.07% 1 0.27% 140 0.13% Gerridae Diapterus peruvianus 1 0.07% 1 0.27% 320 0.30% Ginglymostomatidae cf Ginglymostoma cirratum 1 0.07% 0.00% 0.00% 1500 Haemulidae Anisotremus cf pacifici 1 0.07% 1 0.27% 130 0.12% Haemulidae Haemulopsis elongatus 5 0.33% 4 1.07% 890 0.82% 140-320

Table 7.10: Vampiros 1 and 2 : Fish species list at Macrounit 1 (continued) 206

Family genus Species NISP % MNI % Biomass (gr) % Range Haemulidae Haemulopsis cf leuciscus 1 0.07% 1 0.27% 230 0.21% Haemulidae Haemulopsis leuciscus 5 0.33% 4 1.07% 870 0.80% 120-350 Haemulidae Haemulopsis nitidus 2 0.13% 2 0.53% 300 0.28% 50-250 Haemulidae 23 1.53% 0.00% 0.00% Haemulidae Orthopristis chalceus 76 5.04% 17 4.53% 2700 2.49% 30-240 Haemulidae Pomadasys cf bayanus 1 0.07% 1 0.27% 120 0.11% Haemulidae Pomadasys macracanthus 1 0.07% 1 0.27% 800 0.74% Haemulidae Pomadasys 2 0.13% 1 0.27% 120 0.11% 120-850 Haemulidae Pomadasys panamensis 4 0.27% 4 1.07% 2020 1.86% 220-800 Lutjanidae 1 0.07% 0.00% 0.00% Mugilidae Mugil curema 16 1.06% 3 0.80% 400 0.37% 30-320 Muraenidae Muraena 3 0.20% 0.00% 0.00% 150 Narcinidae Narcine entemedor 1 0.07% 0.00% 0.00% 1400 Paralichthyidae Citharichthys gilberti 1 0.07% 0.00% 0.00% 90 Paralichthyidae Cyclopsetta querna 1 0.07% 0.00% 0.00% 400 Polynemidae 2 0.13% 0.00% 0.00% Polynemidae Polydactylus approximans 32 2.12% 10 2.67% 2465 2.27% 15-400 Polynemidae Polydactylus opercularis 57 3.78% 15 4.00% 4790 4.42% 70-850 Pristigasteridae Ilisha furthii 66 4.38% 13 3.47% 4250 3.92% 80-650 Pristigasteridae Odontognathus panamensis 3 0.20% 0.00% 0.00% 10 Sciaenidae Bairdiella armata 5 0.33% 1 0.27% 100 0.09% 100-130 Sciaenidae Bairdiella ensifera 3 0.20% 3 0.80% 450 0.42% 50-250 Sciaenidae Bairdiella 1 0.07% 1 0.27% 150 0.14% Sciaenidae Cynoscion albus 8 0.53% 6 1.60% 2400 2.21% 150-800 Sciaenidae Cynoscion cf albus 1 0.07% 1 0.27% 1400 1.29% Sciaenidae Cynoscion 9 0.60% 4 1.07% 1900 1.75% 200-2400 Sciaenidae Cynoscion phoxoephalus 1 0.07% 0.00% 0.00% 320 Sciaenidae Cynoscion praedatorius 1 0.07% 1 0.27% 4200 3.88% Sciaenidae Cynoscion squamipinnis 1 0.07% 0.00% 0.00% 450 Sciaenidae Cynoscion cf stolzmanni 1 0.07% 0.00% 0.00% 140 Sciaenidae Cynoscion stolzmanni 9 0.60% 5 1.33% 1390 1.28% 150-650 Sciaenidae Larimus acclivis 2 0.13% 1 0.27% 140 0.13% 70-140

Table 7.10: Vampiros 1 and 2 : Fish species list at Macrounit 1 (continued)

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Family genus Species NISP % MNI % Biomass (gr) % Range Sciaenidae Menticirrhus panamensis 3 0.20% 2 0.53% 300 0.28% 120-500 Sciaenidae Micropogonias altipinnis 4 0.27% 3 0.80% 2450 2.26% 250-1200 Sciaenidae 16 1.06% 1 0.27% 1200 1.11% Sciaenidae Ophioscion 6 0.40% 2 0.53% 150 0.14% 10-140 Sciaenidae Ophioscion typicus 16 1.06% 10 2.67% 1220 1.13% 40-250 Sciaenidae Ophioscion vermicularis 7 0.46% 1 0.27% 80 0.07% 70-150 Sciaenidae Ophioscion cf typicus 2 0.13% 2 0.53% 110 0.10% 30-80 Sciaenidae Ophioscion cf scierus 2 0.13% 2 0.53% 260 0.24% 30-130 Sciaenidae Ophioscion cf vermicularis 1 0.07% 1 0.27% 170 0.16% Sciaenidae Paralonchurus dumerlii 3 0.20% 3 0.80% 1750 1.61% 300-750 Sciaenidae Paralonchurus goodie 1 0.07% 1 0.27% 300 0.28% Sciaenidae Paralonchurus cf dumerlii 1 0.07% 1 0.27% 150 0.14% Sciaenidae Stellifer chrysoleuca 2 0.13% 2 0.53% 320 0.30% 70-250 Sciaenidae Stellifer cf chrysoleuca 1 0.07% 1 0.27% 350 0.32% Sciaenidae Stellifer furthii 2 0.13% 1 0.27% 150 0.14% Sciaenidae Stellifer oscitans 1 0.07% 1 0.27% 20 0.02% Scombridae Scomberomorus sierra 18 1.19% 4 1.07% 2050 1.89% 250-800 Serranidae Rypticus nigripinnis 1 0.07% 0.00% 0.00% 70 Sphyraenidae Sphyraena ensis 17 1.13% 6 1.60% 3350 3.09% 350-850 Sphyrnidae Sphyrna lew ini 2 0.13% 0.00% 0.00% 400-450 Sternopygidae Sternopygus 1 0.07% 0.00% 0.00% 120 Stromateidae Peprilus snyderi 2 0.13% 0.00% 0.00% 200 Stromateidae Peprilus medius 1 0.07% 0.00% 0.00% 250 Tetraodontidae cf Guentheridia formosa 2 0.13% 1 0.27% 360 0.33% Tetraodontidae Guentheridia formosa 44 2.92% 16 4.27% 5380 4.96% 100-600 Tetraodontidae Sphoeroides cf annulatus 2 0.13% 2 0.53% 920 0.85% 120-800 Tetraodontidae Sphoeroides annulatus 32 2.12% 11 2.93% 2480 2.29% 30-500 Total 1507 375 108365

Table 7.10: Vampiros 1 and 2 : Fish species list at Macrounit 1 (continued)

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The pattern in Macrounit 2a is slightly similar (Table 7.11). Here, Pacificmoonfish

(Selene peruviana) represents 8.21% MNI, followed by tete sea-catfish (Ariopsis seemani)

(6.22%), spotted puffer (Guentheridia formosa) (4.88%), yellow bobo (Polydactylus opercularis)

(3.88%), Pacific bumper (Chloroscombrus orqueta) (3.55%), hachet herring (Ilisha furthii)

(3.11%), and Bullseye puffer (Sphoeroides annulatus) (2.89%). In contrast, the percentage of

Estimated biomass indicates that Flap nosed seacatfish (Sciades dowii) contributed a larger proportion of the diet (6%), followed by Kesslers‘ sea catfish (Notarius kessleri) (5.4%), spotted puffer (Guentheridia formosa) (5.3%), tete sea-catfish (Ariopsis seemani) (4.69%), white corvine (Cynoscion albus) (4.57%), Red sea-catfish (Bagre pinnimaculatus) (4.33%), and yellow bobo (Polydactylus opercularis) (4.23%).

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Family genus Specie NISP % MNI % Biomass (gr) % Range Achiridae Achirus scutum 1 0.03% 1 0.11% 150 0.05% Albulidae Albula esuncula 53 1.38% 9 1.00% 4500 1.51% 200-1000 Albulidae Albula 2 0.05% 0.00% 0.00% Albulidae Albula nemoptera 3 0.08% 1 0.11% 500 0.17% Ariidae Ariopsis seemani 234 6.08% 56 6.22% 13965 4.69% 10-1000 Ariidae cf Bagre 1 0.03% 1 0.11% 0.00% Ariidae Bagre 4 0.10% 1 0.11% 600 0.20% 80-600 Ariidae Bagre cf panamensis 1 0.03% 1 0.11% 500 0.17% Ariidae Bagre panamensis 14 0.36% 9 1.00% 3860 1.30% 30-850 Ariidae Bagre cf pinnimaculatus 2 0.05% 0.00% 0.00% 450-800 Ariidae Bagre pinnimaculatus 31 0.80% 13 1.44% 12880 4.33% 180-3500 Ariidae Cathorops cf fuerthii 13 0.34% 8 0.89% 1750 0.59% 30-500 Ariidae Cathorops fuerthii 34 0.88% 11 1.22% 2080 0.70% 40-380 Ariidae Cathorops hypophthalmus 1 0.03% 1 0.11% 300 0.10% Ariidae Cathorops cf multiradiatus 2 0.05% 2 0.22% 360 0.12% Ariidae Cathorops multiradiatus 7 0.18% 3 0.33% 450 0.15% 100-200 Ariidae Cathorops 63 1.64% 16 1.78% 2470 0.83% 40-400 Ariidae Cathorops cf steindachneri 10 0.26% 7 0.78% 1340 0.45% 70-400 Ariidae Cathorops steindachneri 49 1.27% 17 1.89% 3130 1.05% 40-320 Ariidae Cathorops cf tuyra 2 0.05% 2 0.22% 250 0.08% 100-150 Ariidae Cathorops tuyra 4 0.10% 4 0.44% 950 0.32% 200-300 Ariidae 211 5.48% 2 0.22% 850 0.29% 20-1000 Ariidae Notarius cf cookei 1 0.03% 1 0.11% 800 0.27% Ariidae Notarius cookei 3 0.08% 2 0.22% 4400 1.48% 350-4000 Ariidae Notarius cf kessleri 5 0.13% 4 0.44% 1600 0.54% 150-700 Ariidae Notarius kessleri 59 1.53% 25 2.77% 16080 5.40% 140-1500 Ariidae Notarius 69 1.79% 15 1.66% 6500 2.18% 180-1300 Ariidae Notarius troschellii 1 0.03% 1 0.11% 700 0.24% Ariidae Occidentarius platypogon 2 0.05% 1 0.11% 280 0.09% Ariidae cf Sciades dow i 3 0.08% 2 0.22% 1200 0.40% 400-800 Ariidae Sciades dow i 23 0.60% 17 1.89% 17850 6.00% 100-6000 Batrachoididae Batrachoides boulengieri 8 0.21% 4 0.44% 2650 0.89% 250-1000 Table 7.11: Vampiros 1 and 2 : Fish species list at Macrounit 2a

210

Family genus Specie NISP % MNI % Biomass (gr) % Range Batrachoididae Batrachoides pacifici 12 0.31% 5 0.55% 3150 1.06% 250-1200 Belonidae 19 0.49% 3 0.33% 80 0.03% 80 Belonidae Strongylura cf exilis 6 0.16% 0.00% 0.00% 80 Belonidae Strongylura exilis 5 0.13% 1 0.11% 250 0.08% 250-1100 Belonidae Strongylura 5 0.13% 2 0.22% 180 0.06% 80-100 Belonidae Strongylura scapularis 48 1.25% 11 1.22% 2090 0.70% 100-500 Belonidae Tylosurus cocodrilus fodiator 56 1.45% 10 1.11% 2930 0.98% 50-1000 Belonidae Tylosurus imperialis melanotus 1 0.03% 1 0.11% 130 0.04% Belonidae Tylosurus 13 0.34% 5 0.55% 1130 0.38% 40-400 Carangidae Caranx caballus 46 1.19% 11 1.22% 3080 1.03% 100-700 Carangidae Caranx caninus 21 0.55% 11 1.22% 2990 1.00% 50-1000 Carangidae Caranx 9 0.23% 3 0.33% 500 0.17% 50-500 Carangidae Caranx cf caballus 1 0.03% 1 0.11% 250 0.08% Carangidae Caranx cf otrynter 1 0.03% 0.00% 0.00% 200 Carangidae Caranx otrynter 1 0.03% 1 0.11% 150 0.05% Carangidae Caranx vinctus 2 0.05% 2 0.22% 500 0.17% 200-300 Carangidae Chloroscombrus orqueta 96 2.49% 32 3.55% 2140 0.72% 10-150 Carangidae Hemicaranx leucurus 8 0.21% 3 0.33% 700 0.24% 120-300 Carangidae Hemicaranx 1 0.03% 0.00% 0.00% 160 Carangidae Hemicaranx zelotes 6 0.16% 2 0.22% 450 0.15% 150-300 Carangidae 55 1.43% 1 0.11% 5000 1.68% 80-5000 Carangidae Oligoplites altus 5 0.13% 4 0.44% 850 0.29% 80-300 Carangidae Oligoplites 7 0.18% 0.00% 0.00% 80-250 Carangidae Oligoplites cf refulgens 1 0.03% 1 0.11% 100 0.03% Carangidae Oligoplites refulgens 35 0.91% 11 1.22% 1300 0.44% 40-200 Carangidae Oligoplites saurus inornatus 2 0.05% 2 0.22% 250 0.08% 50-200 Carangidae Selar crumenophthalmus 1 0.03% 1 0.11% 230 0.08% Carangidae Selene cf brevoorti 2 0.05% 2 0.22% 400 0.13% 200 Carangidae Selene brevoorti 12 0.31% 7 0.78% 1600 0.54% 100-450 Carangidae Selene 173 4.49% 0.00% 0.00% Carangidae Selene oerstedii 1 0.03% 1 0.11% 180 0.06% Carangidae Selene peruviana 538 13.97% 74 8.21% 8560 2.88% 20-300 Table 7.11: Vampiros 1 and 2 : Fish species list at Macrounit 2a (continued)

211

Family genus Specie NISP % MNI % Biomass (gr) % Range Carangidae Trachinotus kennediy 4 0.10% 2 0.22% 450 0.15% 200-1000 Carangidae Trachinotus 1 0.03% 1 0.11% 1200 0.40% Carangidae Trachinotus paitiensis 3 0.08% 2 0.22% 500 0.17% 150-300 Carcharhinidae Rhizoprionodon longurio 1 0.03% 0.00% 0.00% 350 Carcharhinidae Carcharhinus 1 0.03% 0.00% 0.00% 520 Carcharhinidae cf Negaprion brevirostris 1 0.03% 0.00% 0.00% 2000 Centropomidae Centropomus cf armatus 2 0.05% 1 0.11% 500 0.17% 50-500 Centropomidae Centropomus armatus 4 0.10% 4 0.44% 2150 0.72% 450-700 Centropomidae Centropomus medius 4 0.10% 3 0.33% 2030 0.68% 80-2500 Centropomidae Centropomus 10 0.26% 6 0.67% 1380 0.46% 30-1500 Centropomidae Centropomus cf nigriscens 1 0.03% 1 0.11% 700 0.24% Centropomidae Centropomus nigriscens 3 0.08% 3 0.33% 1350 0.45% 100-1000 Centropomidae Centropomus cf unionensis 2 0.05% 1 0.11% 20 0.01% Centropomidae Centropomus unionensis 8 0.21% 3 0.33% 250 0.08% 50-150 Clupeidae Opisthonema cf libertate 7 0.18% 1 0.11% 200 0.07% 100-200 Clupeidae Opisthonema libertate 119 3.09% 22 2.44% 2800 0.94% 50-250 Cynoglossidae Symphurus elongates 1 0.03% 0.00% 0.00% 10 Eleotridae Eleotris picta 1 0.03% 0.00% 0.00% 80 Eleotridae Dormitator latifrontis 3 0.08% 1 0.11% 250 0.08% 120-250 Elopidae Elops affinis 1 0.03% 0.00% 0.00% 60 Engraulidae Anchoa 1 0.03% 0.00% 0.00% 5 Engraulidae Anchoa panamensis 1 0.03% 1 0.11% 20 0.01% Engraulidae Anchovia macrolepidota 4 0.10% 0.00% 0.00% 20-60 Engraulidae Cetengraulis mysticetus 4 0.10% 1 0.11% 10 0.00% Engraulidae Lycengraulis poeyi 15 0.39% 2 0.22% 150 0.05% 40-150 Ephippidae cf Chaetodipterus zonatus 1 0.03% 0.00% 0.00% 300 Ephippidae Chaetodipterus zonatus 25 0.65% 10 1.11% 4050 1.36% 150-700 Ephippidae Parapsettus panamensis 25 0.65% 3 0.33% 600 0.20% 100-350 Erythrinidae Hoplias microlepsis 1 0.03% 0.00% 0.00% 120 Fistulariidae Fistularia corneta 8 0.21% 2 0.22% 0.00% Gerridae Diapterus peruvianus 9 0.23% 4 0.44% 1500 0.50% 10-400 Gerridae Eucinostomus dow ii 1 0.03% 1 0.11% 90 0.03% Table 7.11: Vampiros 1 and 2 : Fish species list at Macrounit 2a (continued) 212

Family genus Specie NISP % MNI % Biomass (gr) % Range Polynemidae Polydactylus opercularis 193 5.01% 35 3.88% 12595 4.23% 15-800 Pristigasteridae Ilisha furthii 144 3.74% 28 3.11% 9600 3.23% 40-750 Pristigasteridae Odontognathus panamensis 4 0.10% 1 0.11% 10 0.00% Sciaenidae Bairdiella armata 3 0.08% 2 0.22% 300 0.10% 150 Sciaenidae Bairdiella ensifera 1 0.03% 1 0.11% 150 0.05% Sciaenidae Cynoscion cf albus 4 0.10% 0.00% 0.00% 150-3500 Sciaenidae Cynoscion albus 35 0.91% 15 1.66% 13600 4.57% 50-4500 Sciaenidae Cynoscion 52 1.35% 8 0.89% 5150 1.73% 50-3000 Sciaenidae Cynoscion phoxoephalus 1 0.03% 1 0.11% 180 0.06% Sciaenidae Cynoscion cf praedatorius 5 0.13% 3 0.33% 2750 0.92% 350-2500 Sciaenidae Cynoscion praedatorius 22 0.57% 13 1.44% 12180 4.09% 100-3000 Sciaenidae Cynoscion squamipinnis 6 0.16% 4 0.44% 2650 0.89% 250-1500 Sciaenidae Cynoscion cf squamipinnis 1 0.03% 1 0.11% 2000 0.67% Sciaenidae Cynoscion cf stolzmanni 2 0.05% 1 0.11% 70 0.02% 40-70 Sciaenidae Cynoscion stolzmanni 13 0.34% 8 0.89% 1700 0.57% 50-350 Sciaenidae Isopisthus remifer 1 0.03% 1 0.11% 20 0.01% Sciaenidae Larimus pacificus 1 0.03% 1 0.11% 20 0.01% Sciaenidae Larimus acclivis 4 0.10% 2 0.22% 240 0.08% 40-200 Sciaenidae Menticirrhus 1 0.03% 0.00% 0.00% 15 Sciaenidae Menticirrhus cf panamensis 1 0.03% 0.00% 0.00% 60 Sciaenidae Menticirrhus panamensis 3 0.08% 1 0.11% 350 0.12% 60-350 Sciaenidae Menticirrhus elongatus 1 0.03% 1 0.11% 420 0.14% Sciaenidae Micropogonias altipinnis 6 0.16% 3 0.33% 3450 1.16% 70-1800 Sciaenidae 36 0.93% 0.00% 0.00% Sciaenidae Nebris occidentalis 1 0.03% 1 0.11% 320 0.11% Sciaenidae Ophioscion 5 0.13% 1 0.11% 100 0.03% 40-120 Sciaenidae Ophioscion scierus 5 0.13% 3 0.33% 550 0.18% 50-300 Sciaenidae Ophioscion cf scierus 1 0.03% 1 0.11% 150 0.05% Sciaenidae Ophioscion cf typicus 5 0.13% 3 0.33% 310 0.10% 70-200 Sciaenidae Ophioscion typicus 52 1.35% 17 1.89% 2260 0.76% 10-300 Sciaenidae Ophioscion cf vermicularis 1 0.03% 1 0.11% 60 0.02% Sciaenidae Paralonchurus cf dumerlii 4 0.10% 1 0.11% 240 0.08% 50-240

Table 7.11: Vampiros 1 and 2: Fish species list at Macrounit 2a (continued) 213

Family genus Specie NISP % MNI % Biomass (gr) % Range Sciaenidae Paralonchurus dumerlii 10 0.26% 5 0.55% 1280 0.43% 50-450 Sciaenidae Paralonchurus pitersi 1 0.03% 1 0.11% 200 0.07% Sciaenidae Stellifer chrysoleuca 4 0.10% 4 0.44% 910 0.31% 10-400 Sciaenidae Stellifer cf furthii 1 0.03% 0.00% 0.00% 10 Sciaenidae Stellifer furthii 2 0.05% 1 0.11% 30 0.01% Scombridae 1 0.03% 1 0.11% 0.00% Scombridae Scomberomorus sierra 12 0.31% 3 0.33% 850 0.29% 80-700 Serranidae Rypticus nigripinnis 1 0.03% 1 0.11% 80 0.03% Serranidae Rypticus bicolor 1 0.03% 1 0.11% 200 0.07% Sphyraenidae Sphyraena ensis 33 0.86% 9 1.00% 4550 1.53% 150-1000 Sphyrnidae Sphyrna lew ini 2 0.05% 0.00% 0.00% 400-500 Stromateidae Peprilus 1 0.03% 0.00% 0.00% Tetraodontidae cf Guentheridia formosa 6 0.16% 5 0.55% 1900 0.64% 100-650 Tetraodontidae Guentheridia formosa 243 6.31% 44 4.88% 15770 5.30% 30-950 Tetraodontidae 13 0.34% 3 0.33% 1080 0.36% 80-1000 Tetraodontidae Sphoeroides annulatus 168 4.36% 26 2.89% 7770 2.61% 30-700 Tetraodontidae Sphoeroides 3 0.08% 2 0.22% 700 0.24% 80-400 Urotrygonidae Urotrygon asterias 1 0.03% 0.00% 0.00% 200 Total 3851 901 297660

Table 7.11: Vampiros 1 and 2 : Fish species list at Macrounit 2a (continued)

214

Finally (Table 7.12) the pattern observed at Macrounit 2b indicates that

Pacificmoonfish (Selene peruviana) represents 10.76% MNI, followed by spotted puffer

(Guentheridia formosa) (8.35 % ), tete sea- catfish (Ariopsis seemani) (6.86%), green jack (Caranx caballus) (5.57%), yellow bobo (Polydactylus opercularis) (5.57%), blue bobo (Polydcatylus approximans) (3.15%) and Pacificcrevalle jack (Caranx caninus) (2.97%) . In contrast, the percentage of Estimated biomass indicates that spotted puffer fish (Guentheridia formosa) contributes a larger proportion to the diet (8.31%), followed by flap nosed sea-catfish (Sciades dowii)(8%), yellow bobo (Polydactylus opercularis) (5.34%), yellowtail corvina (Cynoscion stolzmanni) (5.22%), tete sea-catfish (Ariopsis seemani) (4.73%) and Pacificmoonfish (Selene peruviana)(4.62%) .

215

Family genus Specie NISP % MNI % Biomass (gr) % Range Albulidae Albula esuncula 8 0.51% 2 0.37% 800 0.45% 300-500 Ariidae Ariopsis seemani 72 4.55% 37 6.86% 8400 4.73% 20-650 Ariidae Bagre 4 0.25% 2 0.37% 900 0.51% 50-500 Ariidae Bagre panamensis 9 0.57% 8 1.48% 4800 2.70% 200-1200 Ariidae Bagre cf pinnimaculatus 1 0.06% 1 0.19% 450 0.25% Ariidae Bagre pinnimaculatus 14 0.89% 6 1.11% 4450 2.51% 400-1200 Ariidae Cathorops cf fuerthii 6 0.38% 5 0.93% 680 0.38% 50-250 Ariidae Cathorops fuerthii 21 1.33% 14 2.60% 2680 1.51% 70-350 Ariidae Cathorops multiradiatus 1 0.06% 1 0.19% 100 0.06% Ariidae Cathorops cf multiradiatus 3 0.19% 2 0.37% 450 0.25% 170-280 Ariidae Cathorops 28 1.77% 13 2.41% 2450 1.38% 50-350 Ariidae Cathorops cf steindachneri 1 0.06% 0.00% 0.00% 100 Ariidae Cathorops steindachneri 45 2.85% 13 2.41% 2300 1.30% 50-400 Ariidae 100 6.33% 6 1.11% 1050 0.59% 50-1000 Ariidae Notarius cf kessleri 3 0.19% 2 0.37% 550 0.31% 150-400 Ariidae Notarius kessleri 16 1.01% 9 1.67% 3300 1.86% 200-1200 Ariidae Notarius lentiginosus 1 0.06% 1 0.19% 400 0.23% Ariidae Notarius 9 0.57% 1 0.19% 800 0.45% 450-800 Ariidae Sciades dow i 21 1.33% 9 1.67% 14200 8.00% 100-5000 Batrachoididae Batrachoides pacifici 2 0.13% 2 0.37% 1000 0.56% 500 Belonidae 2 0.13% 0.00% 0.00% 300 Belonidae Strongylura cf exilis 7 0.44% 1 0.19% 350 0.20% 200-350 Belonidae Strongylura 1 0.06% 1 0.19% 130 0.07% Belonidae Strongylura scapularis 2 0.13% 1 0.19% 140 0.08% Belonidae Tylosurus cocodrilus fodiator 38 2.40% 7 1.30% 2230 1.26% 80-1200 Belonidae Tylosurus imperialis melanotus 1 0.06% 1 0.19% 80 0.05% Belonidae Tylosurus 10 0.63% 4 0.74% 680 0.38% 80-400 Carangidae Caranx cf caballus 1 0.06% 1 0.19% 250 0.14% Carangidae Caranx caballus 70 4.43% 30 5.57% 8150 4.59% 50-600 Carangidae Caranx cf caninus 1 0.06% 0.00% 0.00% 400 Carangidae Caranx caninus 48 3.04% 16 2.97% 5800 3.27% 150-450 Carangidae Caranx 7 0.44% 1 0.19% 0.00% 200-350 Table 7.12: Vampiros 1 and 2: Fish species list at Macrounit 2b 216

Family genus Specie NISP % MNI % Biomass (gr) % Range Carangidae Caranx otrynter 1 0.06% 1 0.19% 200 0.11% Carangidae Caranx vinctus 2 0.13% 1 0.19% 350 0.20% 350-550 Carangidae Chloroscombrus orqueta 27 1.71% 14 2.60% 645 0.36% 15-120 Carangidae Hemicaranx leucurus 11 0.70% 1 0.19% 150 0.08% 100-350 Carangidae 9 0.57% 0.00% 0.00% Carangidae Oligoplites 2 0.13% 0.00% 0.00% 120 Carangidae Oligoplites refulgens 3 0.19% 2 0.37% 280 0.16% 140 Carangidae Oligoplites cf refulgens 1 0.06% 0.00% 0.00% 150 Carangidae Selene 24 1.52% 0.00% 0.00% Carangidae Selene oerstedii 1 0.06% 1 0.19% 270 0.15% Carangidae Selene peruviana 210 13.28% 58 10.76% 8200 4.62% 50-400 Carangidae Trachinotus kennediy 4 0.25% 2 0.37% 1250 0.70% 250-1000 Carangidae Trachinotus paitiensis 3 0.19% 3 0.56% 480 0.27% 70-240 Carcharhinidae Rhizoprionodon longurio 6 0.38% 0.00% 0.00% 350-1000 Centropomidae Centropomus armatus 2 0.13% 1 0.19% 1100 0.62% Centropomidae Centropomus medius 1 0.06% 1 0.19% 150 0.08% Centropomidae Centropomus cf medius 1 0.06% 1 0.19% 150 0.08% Centropomidae Centropomus 8 0.51% 2 0.37% 750 0.42% 20-2000 Centropomidae Centropomus robalito 1 0.06% 0.00% 0.00% 450 Centropomidae Centropomus unionensis 2 0.13% 1 0.19% 600 0.34% 500-600 Centropomidae Centropomus viridis 1 0.06% 1 0.19% 4000 2.25% Clupeidae Opisthonema cf libertate 6 0.38% 0.00% 0.00% 100-150 Clupeidae Opisthonema libertate 70 4.43% 11 2.04% 1320 0.74% 50-250 Clupeidae Opisthonema 2 0.13% 0.00% 0.00% 150 Clupeidae Opisthopterus 1 0.06% 0.00% 0.00% 30 Dasiatidae Dasiatys cf longa 1 0.06% 0.00% 0.00% 1300 Engraulidae Lycengraulis poeyi 3 0.19% 0.00% 0.00% 60 Ephippidae cf Chaetodipterus zonatus 1 0.06% 1 0.19% 350 0.20% Ephippidae Chaetodipterus zonatus 22 1.39% 9 1.67% 3800 2.14% 200-900 Ephippidae 1 0.06% 0.00% 0.00% Ephippidae Parapsettus panamensis 12 0.76% 3 0.56% 750 0.42% 150-250 Fistulariidae Fistularia corneta 3 0.19% 1 0.19% 0.00%

Table 7.12: Vampiros 1 and 2: Fish species list at Macrounit 2b (continued) 217

Family genus Specie NISP % MNI % Biomass (gr) % Range Gerridae Eucinostomus dow ii 1 0.06% 0.00% 0.00% 70 Gerridae Diapterus peruvianus 5 0.32% 4 0.74% 1150 0.65% 200-450 Haemulidae Anisotremus cf pacifici 1 0.06% 0.00% 0.00% Haemulidae Anisotremus dovii 1 0.06% 1 0.19% 500 0.28% Haemulidae Haemulopsis cf elongatus 1 0.06% 1 0.19% 550 0.31% Haemulidae Haemulopsis elongatus 12 0.76% 2 0.37% 950 0.53% 120-750 Haemulidae Haemulopsis cf leuciscus 1 0.06% 1 0.19% 60 0.03% Haemulidae Haemulopsis leuciscus 8 0.51% 5 0.93% 1850 1.04% 70-750 Haemulidae Haemulopsis 1 0.06% 1 0.19% 450 0.25% Haemulidae 7 0.44% 1 0.19% 200 0.11% 150-1400 Haemulidae Orthopristis chalceus 14 0.89% 5 0.93% 770 0.43% 100-230 Haemulidae Pomadasys bayanus 2 0.13% 2 0.37% 1150 0.65% 450-750 Haemulidae Pomadasys cf macracanthus 1 0.06% 1 0.19% 1700 0.96% Haemulidae Pomadasys macracanthus 7 0.44% 5 0.93% 4550 2.56% 150-1800 Haemulidae Pomadasys 4 0.25% 2 0.37% 1950 1.10% 150-1800 Haemulidae Pomadasys panamensis 1 0.06% 1 0.19% 300 0.17% Haemulidae Pomadasys cf panamensis 1 0.06% 1 0.19% 620 0.35% Haemulidae Xenichthys xanti 1 0.06% 1 0.19% 700 0.39% Hemiramphidae Hyporhamphus unifasciatus 2 0.13% 1 0.19% 80 0.05% Hemiramphidae 1 0.06% 1 0.19% 80 0.05% Hemiramphidae Hemiramphus saltator 2 0.13% 0.00% 0.00% 120 Lobotidae Lobotes surinamensis 1 0.06% 1 0.19% 250 0.14% Mugilidae Mugil curema 36 2.28% 9 1.67% 3640 2.05% 50-400 Muraenidae 1 0.06% 0.00% 0.00% 200 Paralichthyidae Citharichthys gilberti 3 0.19% 2 0.37% 120 0.07% 40-80 Polynemidae Polydactylus approximans 34 2.15% 17 3.15% 4350 2.45% 20-450 Polynemidae Polydactylus 1 0.06% 0.00% 0.00% Polynemidae Polydactylus opercularis 90 5.69% 30 5.57% 9480 5.34% 40-700 Pristigasteridae Ilisha furthii 39 2.47% 15 2.78% 5400 3.04% 70-800 Sciaenidae Bairdiella armata 3 0.19% 1 0.19% 70 0.04% 40-70 Sciaenidae Bairdiella ensifera 1 0.06% 1 0.19% 350 0.20% Sciaenidae Cynoscion cf albus 2 0.13% 2 0.37% 1200 0.68% 500-700 Table 7.12: Vampiros 1 and 2: Fish species list at Macrounit 2b (continued)

218

Family genus Specie NISP % MNI % Biomass (gr) % Range Sciaenidae Cynoscion albus 14 0.89% 9 1.67% 7800 4.39% 400-1500 Sciaenidae Cynoscion 18 1.14% 0.00% 0.00% 500-1000 Sciaenidae Cynoscion phoxoephalus 1 0.06% 0.00% 0.00% 530 Sciaenidae Cynoscion praedatorius 5 0.32% 4 0.74% 2470 1.39% 250-1500 Sciaenidae Cynoscion cf praedatorius 1 0.06% 0.00% 0.00% 2000 Sciaenidae Cynoscion cf squamipinnis 1 0.06% 1 0.19% 250 0.14% Sciaenidae Cynoscion squamipinnis 3 0.19% 2 0.37% 650 0.37% 300-350 Sciaenidae Cynoscion cf stolzmanni 1 0.06% 0.00% 0.00% 400 Sciaenidae Cynoscion stolzmanni 10 0.63% 7 1.30% 9270 5.22% 140-4000 Sciaenidae Isopisthus remifer 2 0.13% 0.00% 0.00% 40 Sciaenidae Larimus argenteus 2 0.13% 1 0.19% 70 0.04% Sciaenidae Menticirrhus cf elongatus 1 0.06% 1 0.19% 200 0.11% Sciaenidae Menticirrhus 1 0.06% 1 0.19% 500 0.28% Sciaenidae Menticirrhus elongatus 1 0.06% 1 0.19% 600 0.34% Sciaenidae Menticirrhus panamensis 8 0.51% 1 0.19% 300 0.17% 200-300 Sciaenidae Micropogonias altipinnis 4 0.25% 2 0.37% 950 0.53% 350-600 Sciaenidae 25 1.58% 1 0.19% 200 0.11% 150-550 Sciaenidae Ophioscion cf typicus 4 0.25% 1 0.19% 250 0.14% Sciaenidae Ophioscion typicus 11 0.70% 7 1.30% 680 0.38% 30-250 Sciaenidae Ophioscion 2 0.13% 1 0.19% 60 0.03% 60-180 Sciaenidae Paralonchurus dumerlii 4 0.25% 2 0.37% 900 0.51% 120-500 Sciaenidae Stellifer chrysoleuca 1 0.06% 1 0.19% 200 0.11% Scombridae Scomberomorus sierra 28 1.77% 2 0.37% 850 0.48% 250-1000 Serranidae Epinephelus cf analogus 1 0.06% 1 0.19% 60 0.03% Sphyraenidae Sphyraena ensis 5 0.32% 3 0.56% 750 0.42% 250-1000 Sphyrnidae Sphyrna lew ini 4 0.25% 1 0.19% 500 0.28% 350-500 Tetraodontidae Guentheridia formosa 98 6.20% 45 8.35% 14760 8.31% 100-650 Tetraodontidae 6 0.38% 2 0.37% 0.00% Tetraodontidae Sphoeroides annulatus 37 2.34% 15 2.78% 4330 2.44% 30-650 Tetraodontidae Sphoeroides 1 0.06% 0.00% 0.00% 400 Tetraodontidae Sphoeroides rosemblatti 1 0.06% 1 0.19% 500 0.28% Total 1581 539 181935

Table 7.12: Vampiros 1 and 2 : Fish species list at Macrounit 2b (continued)

219

Figure 7.12: The relative abundance of the species (NISP) at Vampiros 1 and 2 by fish estimated biomass

220

The previous Tables (see range on Tables 7.10 to 7.12) and Figure 7.12 show (on red) that people at Vampiros obtained most of the meat from smaller fish (< 400gr) than (on blue) bigger fish (>400gr). This is similar to behaviour represented in the Early ceramic sites in Central Pacific Panama where early inhabitants preferred smaller and abundant shoaling taxa such as Pacificmoonfish (Selene peruviana), Pacific bumper (Chloroscombrus orqueta) and thread herrings (Opisthonema) which are caught using fine-meshed nets as well as larger specimens available at the estuary (Cooke and Ranere 1999; Cooke 1988, 1992a; Cooke and

Tapia-Rodríguez 1994a).

Natural Transformation Processes and Bones

Several natural processes have been recognized at Vampiros rockshelters which could bias their preservation and subsequently alter our interpretation (Denys 2002; Lyman 1994;

Roberts, et al. 2002). As observed on Figure 7.13, I compared preliminary soil analysis data from both shelters and plotted the evidence of disarticulation, decay and weathering on fish bones. The data suggest different conditions occurring at Vampiros.

Animal transporters had introduced and removed cultural materials, especially at Macrounit -1 as has been presented in a previous chapter. Burrowing activities by wasps, rodents and crabs and the presence of unmodified bat and frog bones have been interpreted as indicative of some disturbance of the deposits and that there may have been some fish bone dispersion (Carvajal Contreras, et al. 2008; Denys 2002).

In terms of decay and articulation, I summarized the completely analyzed material in the following Table (7.13) and Figure 7.10. Articulated cranial bones and caudal vertebrae

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were found mainly in macrounit 2a (162) and the beginning of macrounit 2b (96), few articulated bones were found in macrounit 1 (87). It seems that the rate of deposition was faster in zones with articulated bones than those with less of them. In sum, fewer postdepositional activities affected the macrounit 2 deposits than the macrounit 1 deposits

(Barrett 1997; Denys 2002).

Articulated bones (NISP) Macrounit 1 Macrounit Macrounit 2a 2b Mesh 1/8” 1/16” 1/8” 1/16” 1/8” 1/16” 47 40 111 51 41 55 Total 87 162 96

Table 7.13: Summary articulated bones (NISP) at Vampiros 1 and 2 by mesh and macrounit.

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Figure 7.13: Localization of some natural and cultural bone modifications in Vampiros -1 and 2 stratigraphic profiles

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Evidence of exposure of fish bones on the ground was found at Vampiros-1 (Figure 7.13 and Table 7.14). Some evidence of modifications produced by acid dissolution or exfoliation was observed in the transition between macrounit-2a and 2b. In contrast, hard lumps produced by the concretion or inorganic material were observed more on the surfaces of fish bones at the bottom of macrounit-2b in Vampiros-2. A longitudinal deformation of bone perhaps was caused by trampling, sediment weight, or differential water-logging.

Macrounit 1 Macrounit 2a Macrounit 2b Mesh 1/8‖ 1/16‖ 1/8‖ 1/16‖ 1/8‖ 1/16‖ Concretion 0 0 0 1 15 22 Acid 0 0 0 0 1 0 dissolution/Exfoliation Deformation/Trampling? 3 0 8 2 1 0 Hyperostosis 1 0 0 0 0 0

Table 7.14: Summary natural processes on fish bones (NISP) at Vampiros 1 and 2) by mesh and macrounit.

Hyperostosis is a benign manifestation of normal growth and the characteristic morphology of such hyperostoses have taxonomic value in terms of identification (Bearez

1997). These may take a number of forms: bone swelling of the occipital crest or supraoccipital bone (e.g. Carangidae: Selene oesterdii and Selene brevoorti) and enlargements to occipital crest but also to post-cranial trunk elements (e.g. neural and haemal vertebrae, and anal pterygiophores in Carangidae and Ariidae (Trachinotus and Bagre pinnimaculatus).

Preliminary data of soil chemical conditions indicate that there were slightly more acidic conditions in Vampiros-1 than in Vampiros-2, which could lead to more corrosion and destruction of fish bones (Nicholson 1996: 518). There are not studies that evaluate the density of fish bones on the particular species in Panama. Nicholson´s study is closer to

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Vampiros shelters case since salmon bone structures are not similar to tropical species and it would suggest that Vampiros fish samples were modified by humans. As observed in Table

7.12, there is a noteworthy situation with the greater amount of bones found in Vampiros-1 than Vampiros-2. As well some scales were recovered (5) in macrounit 2a and 2b from

Vampiros-1.

Cultural Processes and Bones

Several alterations on bones are relevant to distinguish cultural origins of bone accumulation.

As with natural processes, cultural modifications could bias the preservation of fish elements, which consequently could alter their interpretation.

Some authors suggest that bones lying on the ground for a long period of time can be trampled by large mammals or humans causing breakage or deformations (Denys 2002).

Other authors suggest that vertebrae deformation; pitting, breakage and rounding are caused by digestive process (Butler and Schroeder 1998). Andrew Jones (1999) also mentions that cultural actions such as trampling affect the state of preservation of fish elements. He suggests after some experiments walking over cod remains and developing and index of robustness that an assemblage of ―recognisable bones consisted of a small number of cranial elements, a few anterior vertebrae and a few extreme caudal centra‖. Also the cleithrum and almost all the trunk vertebrae were absent after a long walk. He said that interpreting remains without the robustness consideration would be mistakenly assumed that as a result

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of decapitating fish and had its tail removed prior to the flesh and trunk vertebrae being transported elsewhere.

However at Vampiros, as observed in Table 7.12, vertebrae deformation was recorded in the transition between macro unit 1 and 2a and 2a and 2b. These deformed vertebrae do not have any other evidence of rounding or pitting. This deformation could result from trampling, water logging or sediment pressure (Figure 7.14a). The relative low numbers of fish bones in Vampiros-2 in comparison to Vampiros-1, it may indicate that the former shelter was used for habitation purposes or smoking fish and the later shelter to cured fish. A great proportion of small bones were recovered on the fine fraction instead of the 1/8‖ mesh of Vampiros-2.

Figure 7.14: Vampiros 1and 2. Preservation and modifications: a. deformed vertebra, b, c &d. different type of scales and e. branchial bones.

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Burning on archaeological bones either may represent cooking activities or discarding activities (Nicholson 1993; Roberts, et al. 2002). Fish bones at Vampiros have been exposed to differential heat as observed in Tables 7.15 and 7.16.

Locus Burnt Total % NISP NISP Vampiros-1 5275 16852 (1/8”mesh) Vampiros-1 21238 34965 (fine mesh) Vampiros-2 888 1160 (1/8”mesh) Vampiros-2 35020 36980 (fine mesh) Total 62421 89957

Table 7.15: (NISP) and percentage of burnt fish remains identified and unidentified by loci and mesh (fine mesh including 1/16‖ and 0.25)

Heating category NISP NISP NISP NISP M1 % M2a % M2b % Grand Total % Complete calcined (white) 25 1.8% 100 3.1% 52 3.9% 177 3.0% Partly calcined (grey) 9 0.6% 14 0.4% 15 1.1% 38 0.6% Partly calcined / partly carbonized 2 0.1% 5 0.2% 30 2.3% 37 0.6% Carbonized (black) 271 19.5% 138 4.3% 259 19.5% 668 11.2% Partly carbonized 3 0.2% 5 0.2% 6 0.5% 14 0.2% Partly carbonized / partly caramelized 6 0.4% 76 2.3% 62 4.7% 144 2.4% Caramelized (red or brown) 111 8.0% 838 25.8% 365 27.5% 1314 22.0% Parly caramelized 17 1.2% 82 2.5% 67 5.1% 166 2.8% Slightly caramelized 44 3.2% 117 3.6% 70 5.3% 231 3.9% Unburnt 900 64.8% 1872 57.7% 400 30.2% 3172 53.2% Grand Total 1388 100.0% 3247 100.0% 1326 100.0% 5961 100.0%

Table 7.16: Differential heat exposure, (NISP) and percentage of burnt fish remains from complete analyzed units by macrounit.

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Table 7.15 suggests in general that 69.3% of the total sample of fish bones at

Vampiros shelters were affected by fire. Specifically, Table 7.15 indicates for the analyzed material that 53.2% of Vampiros‘ sample is unburned, whereas 22 % of fish bones were exposed to 200ºC, 11.2% of Vampiros‘ bones were burned between 300ºC and 500ºC and just a few bones were completely calcined (3.0%).

As observed for both shelters, the soft sediments that correspond to macrounit 1 have few white (1.8%) or black (19.5%) bones and most of them are unburned (64.8%). On the other hand, the deposits at macrounit 2a in both shelters which are characterized by laminar or compact ashes, a mixture of loose sediments, ashes and charcoal present fish bones with several degrees of heat exposure calcined (3.1%), carbonized (4.3%), caramelized

(25.8%) and unburned (57.7%).

For macrounit 2b also, several degrees of heat exposure were observed. This unit, which is characterized by compact ashes and ―greasy‖ sediments, have predominantly fish bones with caramelized (27.5%), black (19.5%) or whitish (3.9%) colors.

These different levels of burning mean if specimens were being cured on drying racks for export I would expect that signs of heat alteration would be for low temperatures

(caramelized) and high temperatures are related to disposal activities (black and whitish).

Relative Representation of Skeletal Elements.

To establish whether fish were decapitated, processed or butchered at Vampiros, I looked at the relative representation of skeletal elements for Macrounit 2. The relative

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distribution of different skeletal elements has been used to understand the exchange of cured fish and site function (Barrett 1997; Barrett, et al. 2008; Belcher 1998; Ceron-Carrasco 1994,

2005; Hoffman, et al. 2000; Zohar 2003; Zohar, et al. 2001a). Several studies assume that if fish were acquired and cured in a processing site such as Vampiros, the site would contain cranial elements and the anterior portion of the vertebral column. Postcranial elements such as the vertebrae and the cleithra would be found at the final destination or consumer site

(Amundsen 2008; Barrett 1997; Hoffman, et al. 2000; Stewart and Gifford-Gonzalez 1994).

Based on the radiocarbon dates, cultural assemblage and the stratigraphic sequence

(Macrounit 2 dated ca. 2200 to 1950 BP), I proposed that both shelters were occupied around the same period; therefore I grouped the 1/8‖ fish sample to conduct element distribution analyses. Due to the speciose nature of the most important food fish families in the tropical Eastern Pacific, it is not practical for the taphonomic analysis at this stage to separate Vampiros samples by species. For this initial assessment, I will group the samples by the most abundant families (Ariidae, Belonidae, Carangidae, Tetraodontidae, Haemulidae,

Pristigasteridae, Polynemidae, Clupeidae and Sciaenidae) that would allow me compare them with Sitio Sierra and Ag-145. Someone could argue that if the species that I group together are a mix large and small species would be an issue to test Zohar's ideas of fish size as the basis for butchering. But as Figure 7.12 show people at Vampiros obtained most of the meat from smaller fish (< 400gr) than bigger fish (>400gr).

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I will include relative the abundance according to anatomic location, and ratio of cranial to postcranial elements and vertebral series for these high-ranking families. The following graphs do not include scales and radials.

As observed in the Ariidae´s Figure (7.15b) cranial bones outnumbered vertebrae.

The presence of caudal and precaudal vertebrae is less in comparison to vertebra complex or

Weberian apparatus (fused 5 or 6 anterior vertebrae)(Figure 7.15c).

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An examination of relative element representation of Ariidae reveals that areas such as the olfactory, occipital, otic, hyoid arch and pectoral girdle are represented in high numbers in comparison to the investing, lateral (maxilla, dentary,premaxilla, dentary, angular, preopercle), opercular, mandibular, pelvic region and caudal skeleton and fins (7.15a).

Figure 7.16: Survival index (SI) for Ariidae fish remains from Macrounit 2

The data from the survival index indicate that the pectoral girdle, occipital, otic(which includes the otoliths), hyoid arch and vertebral column are anatomical regions over-represented in Vampiros sample in comparison to the other ones (Figure 7.16). The underrepresentation of the caudal skeleton, investing, lateral and the opercular region is also

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noteworthy. In Ariidae, the dorsal and pectoral fins are almost equally represented in the sample.

The family Carangidae represents a similar pattern. As shown in graph 7.17b, the cranial skeleton outnumbered the postcranial bones. All anatomical regions are represented at macrounit 2 with the exception of the pelvic girdle and the opercular regions (Figure

7.17a). There is a predominance of precaudal and caudal vertebrae as indicated in the vertebral series graph (Figure 7.17c).

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b c

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The situation described suggests that areas such as the caudal, vertebral column, and olfactory regions are over-represented in comparison to other anatomical regions, the pelvic region being the most underrepresented area (Figure 7.18).

Figure 7.18: Survival index (SI) for Carangidae fish remains from Macrounit 2

For the Clupeidae family, chart 7.16b shows that the cranial skeleton outnumbered the postcranial skeleton.

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The vertebral series indicates that there are more caudal vertebrae than bones from the other sectors (Figure 7.19c). The relative element distribution of this family demonstrates that all the regions are represented with the exception of the occipital, the olfactory, 236

branchial arch and the pelvic girdle (Figure 7.19a). The pectoral girdle and the hyoid arch are more frequent and the pelvic girdle the least frequent. This pattern is confirmed also by the survival index which shows that both the hyoid arch, caudal skeleton, otic, investing, opercular region and the pectoral girdle are overrepresented with values larger than the other anatomical regions (Figure 7.20).

Figure 7.20: Survival index (SI) for Clupeidae fish remains from Macrounit 2

The Haemulidae pattern shows also that cranial bones are slightly more abundant than the postcranial (Figure 7.21b). The precaudal vertebrae are more frequent than the other vertebrae (Figure 7.21c). The relative element distribution (Figure 7.21a) suggests a larger representation of bones from the pectoral girdle, the otic and olfactory region, which is corroborated by the survival index (Figure 7.22). The caudal skeleton and dorsal fin are the most underrepresented areas.

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Figure 7.22: Survival index (SI) for Haemulidae fish remains from Macrounit 2

The Polynemidae family has slightly more postcranial than cranial bones (Figure

7.23b). Thoracic vertebrae are more frequent than the other types of vertebrae (Figure

7.23c). The element distribution chart shows that lateral, hyoid arch, pectoral girdle and the postcranial skeleton are more frequent than the other anatomical regions (Figure 7.23a).

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The survival index chart (Figure 7.24) suggests that the caudal skeleton, vertebral column and hyoid, pectoral girdle and lateral anatomic regions (showing larger values) are over- represented in comparison to other anatomical regions.

Figure 7.24: Survival index (SI) for Polynemidae fish remains from Macrounit 2

The Sciaenidae family also repeats the pattern shown by other families. Graph 7.25b indicates that cranial bones are more frequent than postcranial ones. The vertebral series indicates a significant presence of thoracic vertebrae (Figure 7.25c).

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The relative element distribution suggests that bones from olfatory, occipital lateral, hyoid arch, pectoral girdle , vertebral column and anal fin more frequent than those that

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belong to the investing region, opercular, mandibular , branchial arch , pelvic girdle and dorsal and pectoral fins (Figure 7.25a).

Figure 7.26: Survival index (SI) for Sciaenidae fish remains from Macrounit 2 The survival index also indicates that bones from the anal fin, vertebral column occipital region, lateral region, pectoral girdle and caudal skeleton are more represented than other regions (Figure 7.26).

The pattern observed in the Pristigasteridae family shows that cranial bones are more numerous than postcranial bones (Figure 7.27b). The vertebrae series indicates that precaudal vertebrae are more represented than the other types (Figure 7.27c). The relative

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element distribution of this family suggests that bones from the hyoid arch and the pectoral region are more frequent than other anatomical regions (Figure 7.27a).

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This pattern is also confirmed by the survival index. Graph 7.28 shows that both the pectoral girdle and the hyoid arch are over-represented (showing larger values than other anatomical regions). The branchial arch and the pelvic girdle are poorly represented.

Figure 7.28: Survival index (SI) for Pristigasteridae fish remains from Macrounit 2

The Belonidae family´s bone distribution presents that cranial bones are more numerous than postcranial bones (Figure 7.29b). The vertebrae series indicates that thoracic vertebrae are more represented than the other types (Figure 7.29c).

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The relative element distribution shows that the lateral area is over-represented (Figure

7.29a). The branchial arch, pectoral girdle and pelvic girdle are poorly represented in

Vampiros. This pattern is confirmed by the survival index (7.30). The graph shows that the lateral region is over represented (more than 1) in comparison to the other anatomical regions which are underrepresented (less than 1).

Figure 7.30: Survival index (SI) for Belonidae fish remains from Macrounit 2

Finally, Tetraodontidae´s pattern shows that cranial bones outnumbered the postcranial ones (Figure 7.31b). The vertebral series also indicates a more frequent presence of thoracic vertebrae and precaudal (vertebral column) in comparison to the caudal skeleton

(Figure 7.31c).

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The relative element distribution shows that branchial bones and otoliths are not present in the sample (Figure 7.31a). The lateral region, vertebral column and mandibular region are more frequent than other bones from other anatomical regions. The absence of pelvic girdle is related to the anatomy of this particular family, which for evolutionary reasons does not possess pelvic bones.

Figure 7.32: Survival index (SI) for Tetraodontidae fish remains from Macrounit 2

The survival index chart (Figure 7.32) suggests over-representation of bones from vertebral column, caudal skeleton, hyoid arch, lateral and mandibular region in comparison to other regions.

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Articulated Bones

When fish were butchered either to remove parts (head or tail) or to cure their meat, they may be segmented into convenient portions called butchering units. These also indicate periods of rapid burial after original deposition (Lyman 1994; Reitz and Wing 1999a). As observed in the following table (Table 7.16), most of the articulated bones found came from

Vampiros-1. The majority of the articulations were identified at Macrounit 2a, but none were identified in postholes (―hoyos‖). Most of the fused bones were found in deposits characterized by a mix of soft soil and ash (Vampiros- 1 at subunits 5, 6) but also in compact ashes with burnt bones (Vampiros-1 subunit 29). Most of the articulations in situ from

Vampiros were sections of the vertebral column (ultimate and penultima vertebrae, caudal vertebrae), which suggests that people were disposing of the tails (Figure 7.30b).

Articulated bones 1/8" Vamp-1 Total Vamp-2 Total Grand Total Macrounit 1 Macrounit 2a Macrounit 2b Macrounit 1 Macrounit 2a Macrounit 2b 47 111 41 199 199

Table 7.17: Articulated bones (NISP) at Vampiros 1 and 2 by macrounit.

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Some other articulations suggest that the head was removed as a unit . Three complete catfish neurocrania were found in situ in two macrounits (Figure 7.33a). A vertebral complex articulated to the skull (basioccipital) was also recovered for this family (Figure

7.33h).

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The following units indicate that marine catfish (Ariidae) were gutted removing the head as a unit: in the lateral area , hyoid arch, and pectoral girdle (Table 7.18) .

Family Anatomical area Bones NISP Comments Ariidae NEUROCRANIUM Neurocranium 88 3 individuals

INVESTING-OTIC FRN-SPH 4 2 individuals OCCIPITAL BAS- Vertebral complex 4 2 individuals EPI-SUP 2 1 individual OCCIPITAL-OTIC EPI-PTE-SUP 3 1 individual EPH- CER 2 1 individual EPH- CER-HYH 9 3 individuals CER-HYH 10 5 individuals HYOID ARCH HYO-PRO 2 1 individual PRO-QUA 2 1 individual LATERAL ANG-DEN 2 1 individual COR-CLE 2 1 individual PECTORAL GIRDLE Dorsal spine - Pterygophore dorsal 10 5 individuals SPH-SUP-POS 3 1 individual PECTORAL GIRDLE- OCCIPITAL-OTIC POS-PTE-SUP-SPH 5 1 individual

Table 7.18: Macrounit 2, Family Ariidae: articulated bones by anatomic region

The articulated bones in situ for the Carangidae family suggest that the head was removed intact as a unit, and because of post-depositional processes, it was broken into pieces : an occipital and/or otic region were discarded as a unit , hyoid arch and the olfatory region

(Table 7.19, Figure 7.33e) .

Family Anatomical area Bones NISP Comments OCCIPTAL -OTIC EXO-PTE 2 1 individual OTIC PRT-SPH 2 1 individual HYOID ARCH CER-EPH 2 1 individual Carangidae OLFATORY SET-ETH 2 1 individual

Table 7.19: Macrounit 2, Family Carangidae: articulated bones by anatomic region

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The Sciaenidae, Pristigasteridae, Scombridae and Polynemidae families have a butchering unit on the hyoid arch(epihyal-ceratohyal), which indicates that fish were being gutted (Table 7.20). The Albulidae , Belonidae and Pristigasteridae families have also evidence of fish being decapitated (olfatory region, occiptal-otic region, investing-otic region).

Anatomical Family area Bones NISP Comments Albulidae OLFATORY PRF- PAR 3 1 individual OCCIPITAL- Belonidae OTIC EXO-PTE 2 1 individual PENULTIMA- CAUDAL 2 1 individual Clupeidae CAUDAL CAUDAL 4 1 individual HYOID ARCH EPH-CER 4 2 individuals Polynemidae CAUDAL CAUDAL 2 1individual INVESTING- Pristigasteridae OTIC FRN-PTE 4 1 individual EPH-CER 2 1 individual EPH- CER- HYH 3 1 individual HYOID Sciaenidae ARCH CER-HYH 2 1 individual HYOID Scombridae ARCH EPH-CER 2 1 individual

Table 7.20: Macrounit 2, Families Albulidae, Belonidae, Clupeidae, Polynemidae, Pristigasteridae, Sciaenidae and Scombridae: articulated bones by anatomic region

Finally two butchering units were identified for the Tetraodontidae family at the lateral region (articular-dentary), and olfatory region (ethmoid-supraethmoid) which suggets that people were removing the guts (Figure 7.33f and Table 7.21).

Family Anatomical area Bones NISP Comments LATERAL ANG-DEN 8 4 Individuals Tetraodontidae OLFATORY ETH-SET 6 3 Individuals

Table 7.21: Macrounit 2, Family Tetraodontidae: articulated bones by anatomic region

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As represented in Table 7.20 , articulated vertebrae in situ in the Clupeidae family correspond to caudal and antepenultimate vertebrae. In the following figures, I illustrate the vertebrae represented on Macrounit 2 samples. On the top of each part of the pie the vertebrae position chart is indicated. At the bottom the percentage (NISP%) is indicated.

Families such as Belonidae (Figure 7.34) ,Polynemidae (Figure 7.36), Pristigasteridae,

Haemulidae (Figure 7.35), and Sciaenidae (Figure 7.37), have adjacent caudal vertebrae

(tail) and thoracic (head) vertebrae, which were discarded as separate units after perhaps filleting the specimens. In some families, such as Carangidae ( Figure 7.35), vertebrae from diferent sections were thrown away. The upper section of the vertebral column (thoracic and precaudal vertebrae) were discarded as a unit in families such as Tetraodontidae (Figure

7.37), Ariidae (Figure 7.34), and Belonidae.

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Figure 7.34: Ariidae NISP vertebraeMacrounit 2= 5 Belonidae NISP vertebraeMacrounit 2= 6

Figure 7.35: Carangidae NISP vertebrae Macrounit 2= 398 Haemulidae NISP vertebrae Macrounit 2= 40

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Figure 7.36: Polynemidae NISP vertebrae Macrounit 2= 80 Pristigasteridae NISP vertebrae Macrounit 2= 15

Figure 7.37: Sciaenidae NISP vertebrae Macrounit 2= 15 Tetraodontidae NISP vertebrae Macrounit 2= 87

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Damage

Butchery analysis on fish bones from an archaeological perspective has depended on categorizations (Belcher 1998; Fisher 1995; Zohar 2003) . This is complemented by studies that aim to identify what the marks on the surface of the bones represent (Belcher 1998;

Willis et al 2008). The first step of the analysis was to differentiate beween ancient damage or fractures from recent ones (see Table 7.2).

In the followinf table (Table 7.22), I identified damage on fish bone attributed to ancient damage or butchering practices (identified with the letter ―P‖) , postdepositional and/or taphonomic processes( identified with the letter ―S‖)and indetermined (identified with the letter ―I‖).

The table shows that most of the breaks are distributed on macrounit 2a and 2b. A great proportion of postdepositional cutmarks are found on 1/16‖ mesh material. These postdepostional marks could have been produced during the storage and processing. I kept the the material in bags with sediments.

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1/8" 1/8" Total 1/16" 1/16" Total Grand Total Macrounit 1 Total Macrounit 2a Total Macrounit 2b Total Macrounit 1 Total Macrounit 2a Total Macrounit 2b Total Bone I P S I P S I P S I P S I P S I P S ANG 1 1 1 1 1 1 2 BAM 3 3 3 2 2 2 5 BAS 1 1 1 1 1 2 4 1 1 2 6 7 BR 1 2 3 3 3 CEB 1 1 1 1 1 1 2 CER 1 1 4 1 5 1 1 7 1 1 1 1 2 9 CLE 15 15 15 1 3 4 1 1 3 5 5 5 14 29 COR 1 1 2 2 3 3 RIB 4 4 2 2 6 6 DE 1 5 6 2 2 8 8 DEN 1 1 4 4 1 1 2 7 1 1 1 1 2 3 10 EA 2 2 1 2 3 5 1 1 1 6 ECT 2 2 1 1 3 3 ENT 1 1 0 1 1 EP 2 1 3 3 3 6 3 1 4 4 10 EPB 1 1 1 1 EPH 1 1 1 1 2 1 1 1 3 EPI 2 2 1 1 3 3 UND SPINE 1 1 1 1 2 2 2 1 3 3 7 1 2 2 5 14 16 ETH 2 2 2 2 4 4 FRN 5 5 5 1 1 0 1 6 HYH 1 1 2 1 1 3 1 1 1 4 HYO 5 5 5 2 1 3 4 4 7 12 INT 1 1 1 1 MAX 3 3 1 1 4 1 1 2 1 3 2 2 6 10 MET 1 1 1 1 NT 1 1 1 1 OPE 3 3 3 1 1 1 1 2 5 OTO 1 1 1 1 PAL 1 1 1 1 PAR 1 1 2 2 2 PCM 1 1 2 2 3 3 PETI 1 1 1 1 1 1 3 1 6 7 1 1 2 4 1 1 2 13 16 PM 1 1 1 4 5 2 2 8 1 1 1 1 1 1 3 11 PP 1 1 1 1 PRF 1 1 1 3 1 1 1 3 6 6 PRO 1 1 1 1 PTE 1 1 1 1 1 1 2 PTG 1 1 2 2 1 1 1 3 6 6 QUA 1 1 1 1 3 5 6 6 S 3 3 1 1 4 4 SET 2 2 1 1 3 3 SPC 2 2 1 1 3 3 SPH 1 1 1 1 SUP 2 2 2 2 4 1 1 0 1 5 URO 1 1 1 1 2 1 1 1 3 VERTEBRA 4 4 1 1 1 1 6 2 6 8 2 4 12 18 8 24 32 58 64 VOM 1 1 2 2 3 3 VX 3 1 4 1 1 1 1 6 6 Grand Total 3 18 1 22 2 67 4 73 0 21 1 22 117 1 13 29 43 12 22 58 92 4 30 33 67 202 319

Table 7.22: Vampiros 1 and 2: cur marks by macrounit and mesh size( I= Indeterminate, P= Predepositional, S=Postdepositional)

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Cut marks

As Willis et al. (2008) mention , cut marks on fish are rarely reported because their of their low visibility. They are present not only on diagnostic but on undiagnostic bones. Following

Belcher´s (1998), Fisher (1995), Seetah (2006) and Zohar´s (2003), I observed each bone to identify cut marks, recording its location, number and morphology (Table 7.3). The majority of cut marks tended to be shallow and small striations (cut), impact is like a percussion pit with a circular perforation or depresion , crushing is a splintererd marging of the bone , and chopping mark is smooth entry and fractured exit points .

In this section, I will consider the 1/8‖ samples of Macrounit 2 from both shelters. A total of 44 marks were identified on Ariidae bones. Four of these marks have a postdepostional origin (a different light coloration was observed) and the rest have predepositional origin. Most of the marks are concentrated on the dorsal spine, pectoral spine, ceratohyal and supraoccipital (Table 7.23).

.Bone Predepositional Postdepositional Total CER 4 1 5 CLE 3 3 DE 7 7 DEN 0 1 1 EP 5 1 6 EPH 1 1 UND SPINE 1 1 ETH 1 1 FRN 3 3 HYH 1 1 HYO 2 2 MET 1 1 PETI 2 2 PRO 1 1 SUP 4 1 5 URO 2 2 VX 2 2 Grand Total 40 4 44 Table 7.23: Summary distribution of cut marks at Macrounit 2, Ariidae family

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Bone Mark Number Localization Morphology Direction POSTERIOR CRUSHED 1 /CAUDAL IRREGULAR CHOPPED 2 MID-LATERAL STRAIGHT CER CUT 1 ANTERIOR/DORSAL LINE TRANSVERSAL CHOPPED 2 ANTERIOR/MEDIAL IMPACT CLE CRUSHED 1 MID IRREGULAR CHOPPED 6 MEDIAL-LATERAL IRREGULAR DE CRUSHED 1 MEDIAL-LATERAL IRREGULAR MEDIAL- STRAIGHT- EP CHOPPED 5 POSTERIOR IRREGULAR EPH CHOPPED 1 LATERAL STRAIGHT TRANSVERSAL UND SPINE CHOPPED 1 MEDIAL STRAIGHT ETH CHOPPED? 1 MID STRAIGHT LONGITUDINAL FRN CHOPPED 3 MEDIAL STRAIGHT TRANSVERSAL HYH CHOPPED 1 LATERAL STRAIGHT TRANSVERSAL HYO CUT 1 LATERAL LINE TRANSVERSAL MET CHOPPED 1 MEDIAL STRAIGHT TRANSVERSAL CRUSHED 1 MEDIAL IRREGULAR PETI CHOPPED 1 ANTERIOR STRAIGHT PRO CHOPPED 1 MEDIAL STRAIGHT TRANSVERSAL- SUP CHOPPED 4 MEDIAL-LATERAL STRAIGHT LONGITUDINAL STRAIGHT- URO CHOPPED 2 LATERAL-VENTRAL IMPACT LONGITUDINAL VX CHOPPED 2 LATERAL STRAIGHT LONGITUDINAL

Table 7.24: Distribution of predepostional cut marks at Macrounit 2, Ariidae family

In the Ariidae family, cut marks are concentrated on ceratohyal, supraoccipital, epihyal, cleithrum, pectoral spines and dorsal pterygiophore. They exhibited four patterns: first fine transverse incisions produced perhaps by a stone knife (ceratohyal, hypohyal, hyomandibula, and epyhial, Figure 38d,e). Second; smooth or a splintered marging marks as a result of may be a longitudinal splitting with a polished axe (vertebral complex, urohyal, ethmoid), third a crushing marks observed on pectoral and dorsal spines which produced proximal or medial fragments. And four, lateral and medial cut marks on cranial elements like Belcher (1998) perhaps are the product of splitting the fish to lay it flat either for salting or drying.

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Bone Mark Number Localization Morphology Direction DEN CHOPPED 1 MEDIAL IMPACT HYO CUT 1 MEDIAL LINE TRANSVERSAL PM CHOPPED 1 MEDIAL STRAIGHT Grand Total 3 Table 7.25: Distribution of predepostional cut marks at Macrounit 2,Belonidae family

There are few marks in Belonidae family and all of them have a predepositional origin. They are found in the, premaxilla, dentary and hyomandibular. There are three morphologies: first, fine striations produced by a knife (hyomandibular); second, smooth or percussion pit-like marks (frontal, dentary). Third, cut marks on the medial portion of the cranial skeleton suggest a dried fish processing.

Bone Mark Number Localization Morphology Direction CLE CHOPPED 2 MEDIAL STRAIGHT-IRREGULAR TRANSVERSAL LAT. ETH CHOPPED 1 MEDIAL IRREGULAR TRANSVERSAL SPC CUT 1 LATERAL LINE TRANSVERSAL TRANSVERSAL- VERTEBRA CHOPPED 1 MEDIAL-LATERAL STRAIGHT-LINE LONGITUDINAL Grand Total 5

Table 7.26: Distribution of predepostional cut marks at Macrounit 2,Carangidae family

The Carangidae family presents predepositional marks localized especially in the vertebrae, ceratohyal, lateral ethmoid, supracleithrum, and cleithrum. They describe three patterns: first, transverse fine striations produced by a knife (supracleithrum, precaudal vertebrae); second, smooth marks (lateral ethmoid); and a smooth cut dividing a vertebra in

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a sagittal plane. Most of the cut marks are concentrated on the medial side suggesting that

Pre-Columbian inhabitants were splitting fish either to salt or dry them (Belcher 1998).

Bone Mark Number Localization Morphology Direction BAM CHOPPED 3 MEDIAL STRAIGHT TRANSVERSAL MEDIAL- CLE CHOPPED 5 LATERAL STRAIGHT-IRREGULAR TRANSVERSAL Grand Total 9 Table 7.27: Distribution of predepostional cut marks at Macrounit 2, Haemulidae family

The few predepositional marks observed on the Haemulidae bones were found mainly in the cleithrum and the basypterigium. These are smooth entry and fractured exit points related for splitting the fish for salt and/or drying (Belcher 1998; Ceron-Carrasco 1994).

Bone Mark Number Localization Morphology Direction CLE CHOPPED 1 MEDIAL STRAIGHT TRANSVERSAL DEN CHOPPED 1 MEDIAL STRAIGHT TRANSVERSAL HYO CHOPPED 1 MEDIAL STRAIGHT TRANSVERSAL MAX CHOPPED 1 MEDIAL STRAIGHT TRANSVERSAL Grand Total 4 Table 7.28: Distribution of predepostional cut marks at Macrounit 2, Pristigasteridae family

The few predepositional marks in Pristigasteridae are transverse with a smooth morphology suggesting the use of an axe. They were found in the hyomandibular, cleithrum, dentary, and maxillary. These cut marks suggest people were splitting the fish for salting and/or drying (Belcher 1998)

A total of 8 marks were identified on Sciaenidae bones. Two of these marks have a postdepostional origin (a different light coloration was observed) and the rest have predepositional origin. Most of the marks are concentrated on the maxilla (Table 7.28). 262

Bone Predepositional Postdepositional Total BAS 1 1 DEN 1 1 EA 1 2 3 MAX 2 2 PM 1 1 Grand Total 6 2 8 Table 7.29: Summary distribution of cut marks at Macrounit 2, Sciaenidae family

Bone Mark Number Localization Morphology Direction BAS CHOPPED 1 LATERAL STRAIGHT LONGITUDINAL DEN CHOPPED 1 LATERAL IRREGULAR TRANSVERSAL EA CHOPPED 2 POSTERIOR STRAIGHT TRANSVERSAL MAX CHOPPED 2 DORSAL STRAIGHT TRANSVERSAL PM CHOPPED 1 MEDIAL STRAIGHT TRANSVERSAL Grand Total 7

Table 7.30: Distribution of predepostional cut marks at Macrounit 2,Sciaenidae family

Few cut marks were observed on Sciaenidae bones. These are transverse smooth entry and fractured exit points and were found mainly in maxilla and anal spines. These cut marks suggest that Pre-Columbian inhabitants were splitting the fish and removing the spines (Belcher 1998).

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Bone Mark Number Localization Morphology Direction CLE CHOPPED 1 POSTERIOR STRAIGHT TRANSVERSAL DEN CHOPPED 4 LATERAL-MID-POSTERIOR IMPACT- STRAIGHT LONGITUDINAL FRN CHOPPED 1 MID STRAIGHT TRANSVERSAL MAX CHOPPED 1 MID IRREGULAR TRANSVERSAL PM CHOPPED 5 ANTERIOR-LATERAL-MID LINE-IMPACT TRANSVERSAL SPC CUT 4 LATERAL LINE TRANSVERSAL Grand Total 16 Table 7.31: Distribution of predepostional cut marks at Macrounit 2, Tetraodontidae family

The few predepositional cut marks identified on Tetraodontidae´s bones were found mainly in premaxilla, dentary, frontal, supracleithrum and cleithrum. The cleithrum smooth entry and fractured exit points are produced perhaps from decapitation , and supracleithrum‘s fine striations marks perhaps are result of removing the skin( Figure 38b).

Smooth entry and fractured exit points marks on frontal are perhaps from decapitation.

Percussion pit-like marks are observed on dentary and premaxilla that almost divided the bone longitudinally perhaps to remove the head. Cut marks on the lateral side of the bones suggest that Pre-Columbian inhabitants were removing flesh or skin (Belcher 1998).

Family Bone Mark Number Localization Morphology Direction Albulidae HYO CUT 1 MID LINE TRANSVERSAL Batrachoididae CLE CHOPPED 1 MID IRREGULAR TRANSVERSAL CER CHOPPED 1 LATERAL STRAIGHT TRANSVERSAL Centropomidae CLE CHOPPED 1 MID IRREGULAR TRANSVERSAL HYO CUT 1 VENTRAL LINE TRANSVERSAL Lobotidae OPE CHOPPED 1 LATERAL IRREGULAR TRANSVERSAL ANG CHOPPED 1 ANTERIOR IRREGULAR TRANSVERSAL Lutjanidae DEN CHOPPED 1 ANTERIOR IRREGULAR TRANSVERSAL

Table 7.32: Distribution of predepostional cut marks at Macrounit 2, Albulidae, Batrachoididae, Centropomidae, Lobotidae, Lutjanidae family

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To summarize for the other families, I found striations made by a knife cuts and chopped marks on hyomandibula, cleithrum angular, dentary and opercular perhaps as a result of splitting fish.

Figure 7.38: Fine cut marks: a. HaemulidaeVertebra fine cut mark, b. Tetraodontidae Supracleithrum´s fine cut mark, c.& h. SciaenidaeVertebrae crushed mark, d. & e. Ariidae Ceratohyal cut mark, f. Shark vertebrae chopped mark, g. Sciaenidae Anal spine postdepositional cut mark, notice light color.

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Pattern of Fractures

The analysis of size classes (Table 7.33) demonstrates that of the samples from 1/8‖, 1/16‖ and 0.25mm sieves at Vampiros 17.5% of the bones are highly fragmented, 10.3 % are slightly fragmented and only 34.4% are complete.

Size Class 100% 5% 25% 40% 50% 75% 90% Indet. Grand Total NISP 3191 3 1626 1059 808 813 959 820 9279 % 34.4% 0.0% 17.5% 11.4% 8.7% 8.8% 10.3% 8.8% 100.0%

Table 7.33: General frequency (NISP) and percentage of fragmentation at Vampiros

In macrounits 1, 2a and 2b the bones were complete (ca. 9.53%, 24.07% and 8.40% respectively, see Table 7.34). However highly fragmented bones (between size 5% and 25%) were also identified at macrounits 1, 2a and 2b (ca. 6.91%, 14.83% and 6.48% respectively).

Vampiros 1/8" Size Macrounit 1 Macrounit 2a Macrounit 2b NISP % NISP % NISP % Grand Total 100% 568 9.53% 1435 24.07% 501 8.40% 5% 1 0.02% 25% 346 5.80% 552 9.26% 255 4.28% 40% 66 1.11% 332 5.57% 131 2.20% 50% 107 1.79% 330 5.53% 125 2.10% 75% 113 1.90% 258 4.33% 132 2.21% 90% 170 2.85% 323 5.42% 172 2.88% Indet. 19 0.32% 17 0.29% 10 0.17% Total 1389 23.29% 3248 54.47% 1326 22.24% 5963 100%

Table 7.34: Frequency (NISP) and percentage of fragmentation at Vampiros in completed analyzed samples by macrounit

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Vampiros 1/8" fragments Predepositional Postdepositional Grand Total NISP % NISP % NISP % Total NISP % NISP % NISP % Total NISP % Macrounit 1 Macrounit 2a Macrounit 2b Macrounit 1 Macrounit 2a Macrounit 2b 522 15.09% 1613 46.63% 548 15.84% 2683 299 8.64% 200 5.78% 277 8.01% 776 3459 100.00%

Table 7.35: Frequency (NISP) and percentage of predepositional and postdepositional fractures at Vampiros in completed analyzed samples by macrounit

As shown in Table 7.35, 3459 bones were found in fragmented condition. The majority originated from predepositional events. A few had fractures of postdepositional origin.

Considering the bones differential state of fragmentation in each macrounit, and the different NISP, I address the bones‘ state of fragmentation for macrounits 2a and 2b. The weighted mean index of fragmentation (WMI) will be examined for both macrounits according to the most abundant families: Ariidae, Belonidae, Carangidae, Tetraodontidae,

Haemulidae, Pristigasteridae, Polynemidae, Clupeidae and Sciaenidae (Appendix on CD).For example the WMI for the Olfactory region in Ariidae is calculated as follows:

WMI=[(3·71*100)+(0*5)+(1.6*25)+(2*40)+(1.33*50)+ (1*75)+ (1*90)]/100=7.25%.

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Anatomic region 100% 5% 25% 40% 50% 75% 90% WMI % Olfactory 3.71 0.00 1.67 2.00 1.33 1.00 1.00 7.25 Occipital 6.33 0.00 3.00 2.50 1.50 3.00 3.00 13.78 Otic 15.00 0.00 1.50 1.00 1.00 3.00 2.20 20.51 Investing 4.67 0.00 8.00 0.00 3.00 2.00 0.00 9.67 Lateral 3.80 0.00 2.00 1.00 2.00 1.00 1.33 7.65 Opercular 2.00 0.00 0.00 1.50 2.00 1.33 2.00 6.40 Mandibular 5.33 0.00 3.00 1.00 5.00 1.00 1.50 11.08 Hyoid Arch 11.25 0.00 8.00 6.00 2.00 1.00 2.00 19.20 Branchial Arch 2.00 0.00 2.00 0.00 1.50 3.00 1.67 7.00 Pectoral Girdle 8.75 0.00 10.33 3.00 7.75 2.33 1.67 19.66 Pelvic Girdle 2.00 0.00 1.00 1.00 2.00 1.00 2.00 6.20 Vertebral Column 28.00 0.00 13.00 3.00 2.00 1.00 9.50 42.75 Caudal skeleton 39.00 0.00 0.00 0.00 1.00 4.00 14.00 55.10 Pectoral fin 0.00 1.00 12.00 7.00 4.00 1.00 1.00 9.50 Dorsal fin 20.00 0.00 6.00 2.00 2.50 2.00 4.00 28.65

Table 7.36: Frequency (%) of Ariidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Vampiros (macrounit 2)

In the Ariidae family most of the anatomic areas are highly fragmented (<40%) with the exception of the vertebral column (44.76%) and caudal skeleton (55.10%).

Anatomic region 100% 5% 25% 40% 50% 75% 90% WMI% Olfactory 1.00 0.00 0.00 0.00 0.00 0.00 0.00 1.00 Occipital 2.00 0.00 0.00 0.00 0.00 0.00 0.00 2.00 Otic 1.00 0.00 0.00 0.00 0.00 0.00 0.00 1.00 Investing 1.00 0.00 2.00 1.00 1.00 1.00 3.00 5.85 Lateral 0.00 0.00 3.33 2.00 2.00 1.00 2.00 5.18 Opercular 0.00 0.00 0.00 0.00 1.00 0.00 0.00 0.50 Mandibular 0.00 0.00 0.00 0.00 1.00 0.00 1.00 1.40 Hyoid Arch 1.00 0.00 0.00 0.00 0.00 1.00 4.00 5.35 Branchial Arch 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Pectoral Girdle 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Pelvic Girdle 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Vertebral column 6.67 0.00 0.00 0.00 0.00 1.00 8.00 14.62 Caudal skeleton 4.67 0.00 0.00 0.00 0.00 0.00 1.00 5.57 Table 7.37: Frequency (%) of Belonidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Vampiros (macrounit 2)

In the Belonidae family the WMI values show that most of the anatomic regions are highly fragmented (<40%), however the vertebral column and caudal skeleton are the best preserved.

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Anatomic region 100% 5% 25% 40% 50% 75% 90% WMI% Olfactory 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0 Occipital 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0 Otic 2.00 0.00 0.00 0.00 0.00 0.00 0.00 2 Investing 4.00 0.00 0.00 2.00 1.00 0.00 1.00 6.2 Lateral 0.00 0.00 1.00 1.00 5.00 1.00 0.00 3.9 Opercular 0.00 0.00 1.00 1.00 5.00 1.00 0.00 3.9 Mandibular 1.00 0.00 0.00 0.00 0.00 0.00 0.00 1 Hyoid Arch 2.00 0.00 1.00 2.00 2.00 1.00 0.00 4.8 Branchial Arch 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0 Pectoral Girdle 0.00 0.00 1.00 3.00 2.50 3.00 3.00 7.65 Pelvic Girdle 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0 Vertebral column 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0 Caudal skeleton 26.00 0.00 0.00 0.00 1.00 0.00 1.00 26.5 Table 7.38: Frequency (%) of Clupeidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Vampiros (macrounit 2)

In the Clupeidae family, the occipital, the olfactory, branchial arch, vertebral column and the pelvic girdle are absent regions in the sample which explains the low value of WMI

(0%), and therefore their underrepresentation is not caused by fragmentation. In general, the

WMI values are low; the caudal skeleton being the best preserved region.

Anatomic region 100% 5% 25% 40% 50% 75% 90% WMI% Olfactory 8.75 0.00 2.00 0.00 2.33 0.00 1.00 11.32 Occipital 6.00 0.00 13.00 8.00 5.00 4.00 0.00 17.95 Otic 7.67 0.00 0.00 0.00 0.00 0.00 0.00 7.67 Investing 1.00 0.00 8.00 8.00 10.00 5.00 4.00 18.55 Lateral 1.75 0.00 2.00 3.33 2.50 1.33 1.00 6.73 Opercular 1.00 0.00 1.00 0.00 2.00 1.00 3.00 5.70 Mandibular 3.00 0.00 1.50 1.00 1.33 3.00 2.33 8.79 Hyoid Arch 8.00 0.00 1.50 3.00 5.50 1.33 4.00 16.93 Branchial Arch 3.00 0.00 4.00 3.00 3.00 0.00 0.00 6.70 Pectoral Girdle 1.33 0.00 12.50 15.50 6.50 1.00 2.00 16.46 Pelvic Girdle 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Vertebral column 41.33 0.00 0.00 1.00 3.00 23.00 1.00 61.38 Caudal skeleton 72.00 0.00 0.00 1.00 1.00 4.00 11.67 86.40 Anal fin 4.00 0.00 7.00 6.00 10.00 8.00 3.00 21.85 Dorsal fin 1.00 0.00 0.00 0.00 0.00 0.00 0.00 1.00

Table 7.39: Frequency (%) of Carangidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Vampiros (macrounit 2) 269

Similar to the previous families, for the Carangidae family the WMI values show that most of the anatomic regions are highly fragmented (<40%), however the vertebral column and caudal skeleton are the best preserved.

Anatomic region 100% 5% 25% 40% 50% 75% 90% WMI Olfactory 3.50 0.00 1.00 0.00 1.00 1.00 1.50 6.35 Occipital 2.00 0.00 0.00 0.00 0.00 0.00 1.00 2.90 Otic 3.67 0.00 0.00 0.00 1.00 0.00 1.00 5.07 Investing 4.50 0.00 0.00 0.00 0.00 0.00 1.00 5.40 Lateral 1.00 0.00 1.00 7.00 1.00 0.00 1.00 5.45 Opercular 1.00 0.00 0.00 1.00 1.00 0.00 1.00 2.80 Mandibular 4.00 0.00 0.00 2.00 0.00 0.00 0.00 4.80 Hyoid Arch 1.50 0.00 0.00 1.00 2.00 2.00 0.00 4.40 Branchial Arch 1.50 0.00 1.00 0.00 1.00 0.00 1.00 3.15 Pectoral Girdle 1.00 0.00 1.00 5.00 4.00 2.50 1.50 8.48 Pelvic Girdle 0.00 0.00 5.00 1.00 0.00 0.00 1.00 2.55 Vertebral column 10.00 0.00 0.00 0.00 0.00 1.00 2.00 12.55 Caudal skeleton 13.00 0.00 0.00 0.00 0.00 1.00 1.00 14.65 Dorsal fin 1.00 0.00 0.00 0.00 0.00 0.00 0.00 1.00 Anal fin 1.50 0.00 0.00 0.00 0.00 0.00 0.00 1.50

Table 7.40: Frequency (%) of Haemulidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Vampiros (macrounit 2)

The Haemulidae fracture pattern shows that although the caudal skeleton is underrepresented, their WMI value (14.65%) indicates the best preservation in comparison to the other regions.

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Anatomic region 100% 5% 25% 40% 50% 75% 90% WMI% Olfactory 0.00 0.00 2.00 0.00 0.00 1.00 1.00 2.15 Occipital 1.00 0.00 0.00 0.00 0.00 0.00 0.00 1.00 Otic 1.50 0.00 0.00 0.00 0.00 0.00 0.00 1.50 Investing 1.00 0.00 1.00 0.00 0.00 0.00 0.00 1.25 Lateral 2.00 0.00 1.00 3.00 1.33 2.00 2.00 7.42 Opercular 0.00 0.00 0.00 0.00 1.00 0.00 0.00 0.50 Mandibular 2.50 0.00 0.00 0.00 0.00 0.00 0.00 2.50 Hyoid Arch 4.67 0.00 7.00 3.00 1.00 2.00 3.00 12.32 Branchial Arch 0.00 0.00 0.00 0.00 1.00 0.00 1.00 1.40 Pectoral Girdle 1.00 0.00 0.00 2.00 2.50 1.00 3.00 6.50 Pelvic Girdle 0.00 0.00 2.00 0.00 1.00 0.00 0.00 1.00 Vertebral column 20.00 0.00 0.00 0.00 0.00 2.00 3.00 24.20 Caudal skeleton 17.67 0.00 0.00 0.00 1.00 3.00 7.00 26.72

Table 7.41: Frequency (%) of Polynemidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Vampiros (macrounit 2)

The Polynemidae family shows low WMI values (<40%) on the lateral, hyoid arch, and pectoral girdle, which indicates high fragmentation. The vertebral column (24.20%) and caudal skeleton (26.72%) have slightly higher values that indicate the best preservation in comparison to the other regions.

Anatomic region 100% 5% 25% 40% 50% 75% 90% WMI% Olfactory 0.00 0.00 0.00 1.00 0.00 0.00 0.00 0.40 Occipital 3.00 0.00 0.00 0.00 0.00 0.00 0.00 3.00 Otic 5.50 0.00 0.00 0.00 0.00 0.00 0.00 5.50 Investing 2.00 0.00 0.00 0.00 2.00 0.00 0.00 3.00 Lateral 0.00 0.00 1.00 1.00 1.50 1.00 1.00 3.05 Opercular 0.00 0.00 1.00 0.00 1.00 0.00 0.00 0.75 Mandibular 0.00 0.00 1.00 0.00 0.00 1.50 1.00 2.28 Hyoid Arch 2.75 0.00 1.00 0.00 2.00 1.00 0.00 4.75 Branchial Arch 1.00 0.00 0.00 1.00 0.00 0.00 0.00 1.40 Pectoral Girdle 0.00 0.00 3.00 1.00 3.00 2.33 0.00 4.40 Pelvic Girdle 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Vertebral column 13.00 0.00 0.00 0.00 0.00 0.00 0.00 13.00 Caudal skeleton 29.00 0.00 0.00 0.00 0.00 0.00 0.00 29.00 Dorsal fin 0.00 0.00 0.00 0.00 0.00 0.00 1.00 0.90

Table 7.42: Frequency (%) of Pristigasteridae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Vampiros (macrounit 2)

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In the Pristigasteridae family, the pelvic girdle is underrepresented in the sample, which explains the low value of WMI (0%); therefore, its absence is not cause for fragmentation. In general, the WMI values are low, the caudal skeleton being the best preserved region.

Anatomic region 100% 5% 25% 40% 50% 75% 90% WMI Olfactory 2.00 0.00 2.00 1.00 0.00 2.00 1.00 5.30 Occipital 2.67 0.00 0.00 1.00 0.00 1.00 2.00 5.62 Otic 2.50 0.00 0.00 0.00 0.00 0.00 0.00 2.50 Investing 1.00 0.00 1.00 0.00 0.00 0.00 0.00 1.25 Lateral 1.67 0.00 1.67 1.00 1.50 1.00 1.40 5.24 Opercular 0.00 0.00 0.00 0.00 1.00 1.00 1.00 2.15 Mandibular 2.00 0.00 1.00 1.00 0.00 2.00 1.00 5.05 Hyoid Arch 2.75 0.00 2.00 1.00 2.00 1.00 2.00 7.20 Branchial Arch 4.00 0.00 1.00 1.00 2.00 0.00 0.00 5.65 Pectoral Girdle 1.50 0.00 1.00 8.00 5.00 1.00 1.00 9.10 Pelvic Girdle 1.00 0.00 3.00 0.00 0.00 0.00 0.00 1.75 Vertebral column 8.00 0.00 0.00 0.00 0.00 0.00 4.50 12.05 Caudal skeleton 36.00 0.00 0.00 0.00 1.00 5.00 6.00 45.65 Pectoral fin 1.00 0.00 1.00 0.00 0.00 0.00 1.00 2.15 Anal fin 9.00 0.00 0.00 0.00 0.00 1.00 2.50 12.00

Table 7.43: Frequency (%) of Sciaenidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Vampiros (macrounit 2)

For the Sciaenidae family, the WMI values show that most of the anatomic regions are highly fragmented (<40%); however, the vertebral column is the best preserved.

Anatomic region 100% 5% 25% 40% 50% 75% 90% WMI% Olfactory 1.67 0.00 0.00 1.00 1.00 1.00 2.00 5.12 Occipital 1.67 0.00 0.00 0.00 0.00 0.00 1.00 2.57 Otic 5.00 0.00 0.00 0.00 1.00 0.00 0.00 5.50 Investing 9.00 0.00 1.00 2.00 1.00 1.00 1.00 12.20 Lateral 7.00 0.00 3.00 2.33 1.00 2.00 2.67 13.08 Opercular 1.33 0.00 4.50 6.50 3.33 3.00 2.00 10.78 Mandibular 8.00 0.00 1.00 1.00 1.00 1.50 2.25 12.30 Hyoid Arch 7.00 0.00 2.00 0.00 1.00 4.00 6.50 16.85 Branchial Arch 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Pectoral Girdle 3.00 0.00 7.00 4.50 6.00 1.50 0.00 10.68 Vertebral column 14.67 0.00 0.00 3.00 2.00 2.50 4.67 22.94 Caudal skeleton 13.00 0.00 0.00 0.00 0.00 1.67 4.50 18.30 Table 7.44: Frequency (%) of Tetraodontidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Vampiros (macrounit 2)

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Finally, for the Tetraodontidae family, the WMI values show that most of the anatomic regions are highly fragmented (<40%); however, the vertebral column is the best preserved.

For comparison between the results, I ranked the worst preserved skeletal elements

(<40 %) with NISP larger than 10, in each taxonomic group. While most of the best preserved material in Macrounit 2 corresponds to vertebrae, their common type of fracture is related to the lost of haemal and neural spines or to break off of centra. Wheeler and Jones

(1988:108) consider this break off as a result of post-depositionally trampling or other taphonomic agents. I did not consider this fracture as postdepositional since I did not observe a light coloration.

The remaining bones (see Table 7.45 to 7.48) show several portions due perhaps to choppin on a transverse direction as a result of splitting fish.

Almost Anterior Proximal Lateral Proximal Proximal Distal/medi Distal Bone WMI Rank Proximal art. compl. Cranial Lateral Various p. Centrum centrum end centrum Indet. end/medial medial Medial al end/medial Distal end Distal art. CEB 10.00 7 1 1 1 5 DE 4.85 2 2 5 1 2 1 EP 10.60 9 9 1 7 1 4 4 MET 11.15 10 1 1 5 PRO 8.85 6 1 2 3 1 QUA 7.50 4 5 1 1 VERTEBRA FRAG 10.35 8 31 14 2 2 VX 7.75 5 4 2 2 2 6 3 1 UND: SPINE 5.55 3 1 5 3 1 Ariidae UND. SKULL 2.75 1 11 13

DEN 3.70 1 1 1 5 1 2 Belonidae VERTEBRA FRAG 7.60 2 18 6 1 3

Table 7.45: Weighted mean index (WMI) of highly fragmented bones and portion (NISP) for Ariidae and Belonidae in Vampiros (macrounit 2) 273

Other highly fragmented bones are ceratobranchial, dorsal spine (Figure39a), preopercle, quadrate, metapterygoid, pectoral spine and vertebral complex for Ariidae; dentary, palatine, preopercle, quadrate, urohyal, and coracoid for Carangidae. The

Belonidae´s dentary is highly fragmented. Pectoral and dorsal spines are fragmented when e removed to reduce injuries during the handling of the specimens (Belcher 1998).

Almost Proximal Proximal Proximal Distal/ Distal Bone WMI Rank compl. Ventral Lateral end end/medial medial Medial medial end/medial Distal end Distal art.

COR 6.45 2 6 6 1 2 DEN 7.80 4 1 3 5 2 1 PAL 8.65 5 1 1 1 PRO 6.70 3 2 8 1 1 QUA 9.95 6 5 2 5 1 Carangidae URO 4.30 1 2 1 3 4

Clupeidae COR 8.30 1 5 2 5

CLE 9.30 2 1 1 7 1 1 2 1 Haemulidae PRO 5.25 1 1 6 3

Table 7.46: Weighted mean index (WMI) of highly fragmented bones and portion (NISP) for Carangidae, Clupeidae and Haemulidae in Vampiros (macrounit 2)

The coracoid Clupeidae is highly fragmented as well as the cleithrum and preopercle for Haemulidae.

Almost Proximal Lateral Proximal Proximal Distal Bone WMI Rank Proximal art. compl. Caudal Cranial end centrum end/medial medial Medial end/medial Distal end

ANG 7.80 1 2 4 1 HYO 14.60 2 2 5 2 3 3 2 2 Polynemidae Pristigasteridae CLE 5.05 1 1 1 1 5 3

CEB 7.95 3 4 CLE 7.75 2 1 8 2 2 1 Sciaenidae HYO 7.25 1 2 3 1 3 1

Table 7.47: Weighted mean index (WMI) of highly fragmented bones and portion (NISP) for Polynemidae, Pristigasteridae and Sciaenidae in Vampiros (macrounit 2)

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Low WMI values show high fragmentation on articular and hyomandibular for

Polynemidae; cleithrum for Pristigasteridae. Other highly fragmented bones are ceratobranchial, hyomandibular, and cleithrum, for Sciaenidae and first branchiostegal ray, ceratohyal, articular, cleithrum, premaxilla, palatine, quadrate, indeterminate spine, preopercle and frontal for Tetraodontidae.

Almost Proximal Proximal Distal Bone WMI Rank Proximal art. compl. Lateral end end/medial Medial end/medial Distal end

ANG 9.40 2 1 2 BR 10.20 5 4 1 1 1 1 8 CER 13.10 8 4 3 3 CLE 9.60 3 1 1 6 5 8 FRN 12.50 7 1 2 2 1 PAL 9.65 4 2 2 3 PM 14.00 10 2 1 1 1 1 PRO 9.10 1 4 1 2 7 1 1 QUA 13.65 9 2 1 2 Tetraodontidae UND. SPINE 10.60 6 2 2 Table 7.48: Weighted mean index (WMI) of highly fragmented bones and portion (NISP) for Tetraodontidae in Vampiros (macrounit 2)

Typical fractures observed for all families are on the cleithrum which represented several portions (see Table 7.49 and Figure 39b,d). This is due to perhaps transverse cuts either to facilitate salt absorption or to remove the head (Belcher 1998; Zohar and Cooke

1997).

Other broken areas above like the dentaries, opercle and quadrate are perharps damaged due to their vicinity to premaxilla during the splitting process or bones assocated to pectoral girdle and hyoid arch are damaged during the evisceration process.

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Zone Ariidae Carangidae Clupeidae Haemulidae Polynemidae Pristigasteridae Sciaenidae Tetraodontidae Grand Total Complete 6 1 1 8 Almost complete 3 1 1 5 Cranial 1 1 Lateral 1 2 3 Proximal end 19 17 1 1 1 1 40 Proximal end/medial 4 9 7 3 1 8 1 33 Proximal medial 6 1 1 1 9 Medial 11 15 3 1 1 5 2 6 44 Distal/Medial 2 2 1 1 6 Distal end/medial 5 2 4 2 2 5 20 Distal end 2 1 1 3 1 8 16 Grand Total 50 58 9 15 7 11 14 21 185

Table 7.49: Types of fractures observed on the cleithrum on the most abundant families at Vampiros.

Figure 7.39: Chopped marks and fractures: a. Fractured dorsal spine (Ariidae), b. & d. transverse crushed fractures in cleithrum (Ariidae), c. longitudinal fracture in ethmoid (Ariidae), e. transversal fracture in meptaterygoid (Ariidae), f. longitudinal fracture premaxilla (Tetraodontidae), g. Chopped coracoid (Ariidae), h. Chopped frontal (Tetraodontidae), i. Chopped supraoccipital (Ariidae).

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Summary

At the beginning of the occupation, the marine fishing environment surrounding the

Vampiros shelters was characterized by clear-water fish species, found at the outer margin of the mixing plume. At the time of the shelters‘ abandonment, some fish species that prefer turbid estuarine conditions increased. It is possible that these shelters were occupied during the dry season because of the presence of anchovies and spadefish. Similar to Early ceramic sites in Central Pacific Panama, Vampiros inhabitants preferred smaller and abundant shoaling taxa such as Pacific moonfish (Selene peruviana), which were probably fished using fine-meshed nets, and to a lesser extent, larger specimens available at the estuary. Contrary to expectations, Vampiros shows an abundance of puffer fish bones, which suggests their consumption as food.

Natural processes, such as encrustation and animal burrowing, and the activities of the peoples who occupied the shelters, such as trampling, cooking, butchering, curing fish and digging postholes, have had a post-depositional impact on the integrity and preservation of fish remains. Although a cautious interpretation is necessary, butchering could explain the absence and state of preservation of the assemblage, but as I mentioned previously,

Vampiros shelters show intense human activity ,especially in macrounit 2 (i.e. eating, trampling, digging, burning) as well as the impact of natural factors, especially in macrounit

1. In general, the few articulated fish specimens and postdepositional marks indicate a rapid burial, primary deposition and integrity of the deposits. It is not possible to determine if fishbone variations in color are related to cooking or curing fish. Perhaps, bones with the lowest temperature (brown-caramelized) were involved in curing activities, whereas burned bone (black-white) represent discarding activities into hearths.

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To understand if Pre-Columbian inhabitants processed fish in Vampiros, the abundance of elements (%NISP) was organized in a graphic manner, grouping the macrounits. In general, the preponderance of the hyoid arch, branchial apparatus and pectoral girdle in comparison to other regions suggests that these bones were discarded as a result of gutting activities and fractures are associated to prepare fish for salting and drying

(Belcher 1998; Tourunen 2008; Zohar and Cooke 1997).

Figure 7.40: Ariidae specimens from Vampiros Macrounit 2. NISP= 835

In brief, the pattern observed for the Ariidae family suggests that at Vampiros more cranial bones than postcranial bones which mean that the head was cut off and discarded here but vertebrae were removed elsewhere attached to the flesh. The relative distribution of elements indicates that people were removing the gills, fins and tails. The cut-mark 278

distributions and animal suggests that Pre-Columbian inhabitants were perhaps splitting down the midline and drying fish. There are no missing bones for small fish. In fact, three articulated skulls were found and fragments from the vertebral complex, which means that the smaller fish were gutted keeping the head intact. The damage was found on the cleithrum, coracoid, premaxilla, quadrate, metapterygoid, epihyal, preopercle, interopercle and urohyal. There is a high frequency of first dorsal spines are consistent to remove them to avoid injuries. That pattern suggests that people removed the gills and guts, and afterward chopped the fish down the miline, leaving marks in the medial side of the bones.

Figure 7.41: Carangidae specimens from Vampiros Macrounit 2. NISP= 1065

The family Carangidae describes a similar pattern. People were discarding more cranial bones than postcranial ones. Perhaps some cranial bones are more susceptible to damage than

279

others (see better cleithra´s preservation in green) despite of fragmentation as a result of splitting the head. All anatomical regions are represented at Vampiros shelters with the exception of the pelvic girdle which suggest that fish were gutted ventraly, and there is a predominance of precaudal vertebrae. As noted above this distribution and the cut marks suggest that Vampiros inhabitants were fishing smaller animals. There are no missing bones.

Damage on cranial and postcranial bones are related either to cutting (cleithrum) or to their fragility and density (supraoccipital and frontal), as indicated by their fragmented condition.

In this family scales were identified belonging to the genus Caranx. In summary, people scaled the small specimens and removed the gills, guts and fins.

Figure 7.42: Clupeidae specimens from Vampiros Macrounit 2. NISP= 84

For the Clupeidae family a different pattern is observed. These are smaller fish; however, of the cranial bones represented, there is a notable absence of bones from the

280

olfactory and occipital region. The pectoral girdle (cleithrum and coracoid), the hyoid arch and caudal vertebrae are more frequent than the pelvic girdle. The absence of the pelvic girdle is related perhaps to gutting the fish ventrally. The distribution of the vertebrae suggests that the whole anterior section of the animal is missing in Vampiros deposits and that the Pre-Columbian inhabitants were perhaps removing the guts and transporting these anteriore vertebrae with the ―complete‖ (see diagram on white) skull to a consumer site.

Figure 7.43: Haemulidae specimens from Vampiros Macrounit 2. NISP= 201

As described in a previous section, the Haemulidae pattern shows that cranial bones are more abundant than the postcranial, the best preserved being the cleithra (purple) and preopercles (green). The precaudal vertebrae are more frequent than the other vertebrae

(yellow). The relative element distribution suggests a larger representation of bones from the pectoral girdle (cleithrum) which indicates that people were gutting the fish. This distribution and the cut marks information suggests that people were preparing especially small

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specimens (see estimated biomass), removing heads and splitting them for salting and/or drying.

Figure 7.44: Polynemidae specimens from Vampiros Macrounit 2. NISP= 291

The Polynemidae family has more postcranial than cranial bones. Thoracic vertebrae

(in blue) are more frequent than the other type of vertebrae. In fact, the whole mid-section of the specimens is missing. The element distribution suggests that the vertebral column, lateral, hyoid arch and pectoral girdle are more frequent, the angular, hyomandible (in purple indicated >5%) and thoracic vertebrate being the best preserved. This distribution indicates that Pre-Columbian inhabitants were gutting fish. The damage was found on the cleithrum, hyomandible, and preopercle. This distribution suggests that people were not removing the head.

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Figure 7.45: Sciaenidae specimens from Vampiros Macrounit 2. NISP= 320

The Sciaenidae family repeats the pattern shown by other families There are more cranial than postcranial bones. The relative element distribution suggests that bones from investing, opercular, mandibular, and branchial arch regions are more frequent than those that belong to the pelvic girdle and fins, the preopercle, hyomandible and brachostegal ray being the best preserved (in blue). Also bones from the anal fin are more frequently represented than other regions. People were removing gills, guts and fins and disposing them at Vampiros, and the medial cut marks on dentary, premaxilla, basioccipital and spines suggest that they were perhaps people were splitting the head and removing the spines to avoid injuries .

283

Figure 7.46: Pristigasteridae specimens from Vampiros Macrounit 2. NISP= 140

The pattern observed in the Pristerigaridae family shows that thoracic and precaudal vertebrae are more frequently represented than the other types, and the relative element distribution of this family suggests that bones from the hyoid arch (hyomandibular in orange) and the pectoral region (cleithra in purple)are more frequent and survived better than other anatomical regions. This implies again that people were gutting fish. The fragility of some bones is attested by their fragmented condition and damage as seen on the maxilla, hyomandible and frontal. Finally, these fractures and medial cut marks suggest people were splitting for salting and/or drying fish.

284

Figure 7.47: Belonidae specimens from Vampiros Macrounit 2. NISP= 130

Briefly, the pattern observed for the Belonidae family suggests that the Pre-

Columbian inhabitants were disposing of more cranial bones, especially the bones from the lateral area. The element and cut-mark distributions suggest that they were removing the dentary and premaxilla.

285

Figure 7.48: Tetraodontidae specimens from Vampiros Macrounit 2. NISP= 442

To sum up, the pattern observed for the Tetraodontidae family suggests that at

Vampiros cranial bones are more abundant than the postcranial ones. The relative distribution of elements indicates that bones from thoracic vertebrae and the lateral and mandibular regions are over-represented, not only due to butchering practices but also due to differences in their preservation (see premaxilla and opercle in orange, dentary in blue).

The element and cut-mark distributions suggests that Pre-Columbian inhabitants were perhaps removing the skin as striations are found on the supracleithrum, guts (fractures on cleithra and 1st branchiostegal ray), tail (distribution of caudal vertebrae) and head (fractures on frontal) of the animal and popping-out two dorsal fillets.

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Chapter 8

Pre-Columbian Fishing at other sites on the Santa Maria´s Watershed

Introduction

The purpose of this chapter is to provide comparative data to investigate the exchange and production of cured fish at Cueva de los Vampiros. I have suggested that both shelters at

Vampiros might have served as a specialised station to cure and prepare marine fish to distribute to other inland sites up the Santa Maria River Valley (Carvajal Contreras, et al.

2008; Cooke and Jiménez 2008; Cooke and Ranere 1999; Cooke 1988, 1992a). During the

Proyecto Santa Maria (PSM) large village sites were identified along the Santa Maria River.

One of these large sites was Sitio Sierra (Ag-3), which covers approximately 45 ha with cultural remains consistent with a large village and with discrete burial grounds (Weiland

1984). Additionally, this site has provided the largest sample of fish remains of any of the sites along the Santa Maria River. A second site in the Santa María drainage from which fish samples have been identified, is AG-125. This site is one of four oval mounds that were located in 1982 during transect and purposive surveys of the high tidal flats that stretch northwards and eastwards of the Vampiros shelters. They probably belonged to a single cluster of dwellings arranged along an ancient beach. Here I will compare the two faunal assemblages from these sites on the Santa Maria watershed: Sitio Sierra and AG-125, not only because the nature of animal remains might indicate the type of site (processing site vs.

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habitation site) but also because these sites and Vampiros might have similar butchering patterns based on a shared cultural tradition.

Background of the Comparative samples

Sitio Sierra´s sample contained large number of vertebral remains that were excavated and recovered by Cooke in 1971, 1973 and 1975 who published these data in several manuscripts

(Cooke 1993; Cooke, et al. 2008; Cooke and Ranere 1984, 1992a, b; Cooke and Ranere 1999;

Cooke 1979, 1984, 1988, 1992a; Isaza-Aizpurua 1993). Cooke (1993; Cooke and Ranere

1999; 1992a) has presented the taxonomic evaluation of these fish remains, which were recovered from a 4.5x1.5 m, test pit, 0.5m deep, using a 3.2 mm (1/8‖) standard commercial wire mesh. Isaza (1993) called this deposit ―Context B-3‖. It consisted of a circular feature with a diameter of ca 7 m. Although this deposit has no radiocarbon-dates; it lies on top of a probable dwelling and adjacent refuse lenses, associated with charcoal dates of 2015 ± 80

BP (cal BC 190-cal AD 155) and 1975 ± 80 BP (cal BC 170-cal AD 230) (Cooke and Ranere

1984; Isaza-Aizpurua 1993). Additionally, the pottery associated with this context, where the fish bones are embedded, belongs to the types and varieties of the Artistide ceramic group

(Isaza-Aizpurua 1993; Ladd 1964), which ranges between cal AD 250 to 550.

It is likely that this refuse pile was deposited during this period. Thus it overlaps in time with the later stages of the fishing camp at Vampiros.

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AG-125 was a 49 X 38m mound occupied after AD 1000. Excavations revealed floors of structures and refuse lenses, mostly of Crassostrea shells. A test pit (1 x 0.5 m) dug in AG-

125 in 1982 reached sterile beach deposits. The fish sample that Cooke identified, which has not been formally published, represents 5 kg of soil taken from the fifth natural layer lying on sterile soil and was also recovered using 3.2 mm mesh (1/8mm). A charcoal sample of

825 ± 60 BP (cal AD 1040-1285) (GX-25700-LS) was obtained from this soil (Carvajal

Contreras, et al. 2008; Cooke and Ranere 1984; Weiland 1984)

In an earlier paper, I proposed the hypothesis that Ag-125 may has replaced the

Vampiros shelters as the primary fishing settlement at the mouth of the Santa María River, as coastal progradation would have left Vampiros an inconvenient distance from the sea to exploit marine resources (Carvajal Contreras, et al. 2008; Cooke and Ranere 1984; Weiland

1984).

Methodology

Since these samples are taxonomically analyzed, I recorded information about the portion of the skeletal elements, percentage of completeness and origin of fragmentation (see chapter seven). Additionally, the location, number and morphology of the cut marks were also observed. Similar to Vampiros samples, encrustation, articulated bones and acid dissolution as well as cultural processes of heating and trampling of fish bones were registered in both samples (Belcher 1998; Lyman 1994; Zohar 2003; Zohar and Cooke 1997; Zohar, et al.

2001). 289

I calculated the relative representation of skeletal elements based on NISP and MAU values of the most abundant families: Ariidae, Carangidae, Clupeidae, Haemulidae, Tetraodontidae, and Sciaenidae. Similar to Vampiros samples, I included in the analysis the ratio between cranial and post cranial bones, body part frequency, the survival index (SI), and the weighted mean index (WMI) of fragmentation for both samples (Belcher 1998; Lyman 1994; Reitz and

Wing 1999a; Wheeler 1989; Zohar 2003; Zohar and Cooke 1997; Zohar, et al 2001).

Sitio Sierra´s Sample (AG-3)

The 15,549 fish skeletal elements recovered in this sample were identified to 13 families, 34 genera and 63 species; I did not analyze or include freshwater fish skeletal elements (see

Table 8.1 for detailed information). In terms of MNI, the most abundant families in order of importance were Clupeidae (23.3%), Haemulidae (22.1%), Carangidae (17.7%), Ariidae

(16%) and Sciaenidae (4.9%). Noteworthy in particular are the genera Opisthonema (23%),

Orthopristis (17%), Selene (10.8%), Notarius (7.2%) and Pomadasys (3.9%). Specifically the most abundant species were Opisthonema libertate (23%), Orthopristis chalceus (17%),

Selene peruviana (9.8%), Sciades dowii (3.5%) and Dormitator latifrontis (3%) (Cooke and

Jiménez 2004; Cooke and Ranere 1999).

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Estimated Estimated Body Genus and Species NISP % MNI % Body Mass % Genus and Species NISP % MNI % Mass % Elamosbranchs 108 0.66% 0.00% 0.00% Trachiniotus kennedyi 2 0.01% 1 0.13% 3 1.18% Elamosbranch, unidentified species35 0.21% 2 0.27% ? ? CENTROPOMIDAE CARCHARHINIDAE Centropomus 21 0.13% 0.00% 0.00% Carcharhinus cf altinus 2 0.01% 1 0.13% 1 0.39% C. medius 3 0.02% 1 0.13% 0.075 0.03% C.leucas 6 0.04% 2 0.27% 11 4.34% C. cf medius 2 0.01% 1 0.13% 0.8 0.32% C.limbatus 1 0.01% 1 0.13% ? ? C. nigrescens/ viridis 47 0.29% 9 1.21% 14.55 5.74% Rhizoprioniodon longurio 33 0.20% 2 0.27% 1 0.39% C. robalito 1 0.01% 1 0.13% 0.075 0.03% Ray 1 0.01% 0.00% 0.00% CHARACIFORMES 2 0.01% 0.00% 0.00% DASYATIDAE CTENOLUCIIDAE Dasyatis cf longus 1 0.01% 1 0.13% ? ? Ctenolucius hujeta 17 0.10% 3 0.40% 0.225 0.09% MYLIOBATIDAE CURIMATIDAE Aetobatus narinari 4 0.02% 1 0.13% 3 1.18% Curimata magdalenae 26 0.16% 7 0.94% 0.355 0.14% SPHYRNIDAE ERYTHRINIDAE Sphyrna 1 0.01% 1 0.13% ? Hoplias sp 2090 12.81% 50 6.74% 15.55 6.14% cf Sphyrna 1 0.01% 1 0.13% >10 ? CICHLIDAE Sphyrna cf tiburo 19 0.12% 1 0.13% 1 0.39% Aequidens coeruleopunctatus 26 0.16% 9 1.21% 0.4 0.16% PRISTIDAE CLUPEIFORMES 105 0.64% 0.00% 0.00% Pristis 3 0.02% 1 0.13% ? ? CLUPEIDAE UROLOPHIDAE 1 0.01% 1 0.13% ? ? Opisthonema cf libertate 3263 20.00% 119 16.04% 11.9 4.70% Teleosts ENGRAULIDAE Teleost, unid species 1 0.01% 1 0.13% 0.1 0.04% cf Anchoa 3 0.02% 1 0.13% 0.1 0.04% SILURIFORMES 52 0.32% 0.00% 0.00% PRISTIGASTERIIDAE ARIIDAE 1529 9.37% 0.00% 0.00% Ilisha furthii 76 0.47% 5 0.67% 1.95 0.77% Notarius 85 0.52% 1 0.13% 0.025 0.01% EPHIPPIDAE Cathorops dasycephalus 2 0.01% 1 0.13% 0.15 0.06% Parapsettus panamensis 1 0.01% 1 0.13% 0.15 0.06% Cathorops cf dasycephalus 1 0.01% 0.00% 0.00% GERREIDAE Notarius biffi 89 0.55% 10 1.35% 9.5 3.75% cf Diapterus peruvianus 1 0.01% 1 0.13% 0.15 0.06% Notarius cf biffi 5 0.03% 0.00% 0.00% GOBIIDAE/ELEOTRIDAE Notarius kessleri 111 0.68% 13 1.75% 9.3 3.67% Dormitator latifrontis 223 1.37% 15 2.02% 3.575 1.41% Notarius cf kessleri 11 0.07% 0.00% 0.00% Eleotris picta 18 0.11% 4 0.54% 1.225 0.48% Notarius osculus 14 0.09% 3 0.40% 1.5 0.59% Gobiomorus maculatus 2 0.01% 1 0.13% 0.075 0.03% Notarius cf osculus 1 0.01% 0.00% 0.00% cf G. maculatus 2 0.01% 1 0.13% 0.15 0.06% Gn sftep a platypogon 1 0.01% 1 0.13% 0.9 0.36% HAEMULIDAE 467 2.86% 0.00% 0.00% Gn sftep acf platypogon 1 0.01% 0.00% 0.00% Anisotremus 3 0.02% 0.00% 0.00% Ariopsis seemani 80 0.49% 8 1.08% 2.05 0.81% Anisotremus dovii 16 0.10% 2 0.27% 0.9 0.36% cf Ariopsis seemani 5 0.03% 0.00% 0.00% A. pacifici 13 0.08% 4 0.54% 1.45 0.57% Bagre 12 0.07% 0.00% 0.00% Orthopristis chalceus 2967 18.18% 87 11.73% 13.25 5.23% B.panamensis 48 0.29% 6 0.81% 2.6 1.03% Pomadasys 6 0.04% 0.00% 0.00% B. pinimaculatus 35 0.21% 5 0.67% 6.825 2.69% Pomadasys bayanus 7 0.04% 2 0.27% 1.8 0.71% Cathorops 114 0.70% 0.00% 0.00% P. cf bayanus 1 0.01% 1 0.13% 0.2 0.08%

Table 8.1: Sitio Sierra: Fish species list 291

Estimated Estimated Body Genus and Species NISP % MNI % Body Mass % Genus and Species NISP % MNI % Mass % Cathorops (not fuerthii or hypophthalmus)1 0.01% 0.00% 0.00% P. macracanthus 32 0.20% 6 0.81% 4.15 1.64% Cathorops fuerthii 13 0.08% 3 0.40% 0.85 0.34% P. panamensis 2 0.01% 2 0.27% 1.2 0.47% Cathorops cf fuerthii 1 0.01% 1 0.13% 0.03 0.01% P. cf panamensis 1 0.01% 1 0.13% 0.5 0.20% C. fuerthii or hypophthalmus 1 0.01% 0.00% 0.00% Haemulopsis 19 0.12% 2 0.27% 0.45 0.18% C. steindachneri 8 0.05% 4 0.54% 0.575 0.23% H. elongatus 3 0.02% 2 0.27% 0.35 0.14% C. cf steindachneri 3 0.02% 0.00% 0.00% H. cf elongatus 1 0.01% 0.00% 0.00% C. fuerthii or tuyra 1 0.01% 0.00% ? cf H. elongatus 3 0.02% 0.00% 0.00% C. multiradiatus 5 0.03% 2 0.27% 0.35 0.14% H. leuciscus 6 0.04% 2 0.27% 0.75 0.30% C. cf multiradiatus 1 0.01% 0.00% 0.00% H. cf leuciscus 2 0.01% 1 0.13% 0.1 0.04% C. tuyra 20 0.12% 4 0.54% 1.15 0.45% H. cf nitidus 1 0.01% 1 0.13% 0.25 0.10% C. cf tuyra 4 0.02% 0.00% 0.00% LOBOTIDAE Notarius troschelii 3 0.02% 2 0.27% 0.8 0.32% Lobotes pacificus 8 0.05% 3 0.40% 7.25 2.86% Sciades dowii 333 2.04% 18 2.43% 35.45 13.99% LUTJANIDAE cf S. dowii 8 0.05% 0.00% 0.00% Lutjanus cf colorado 1 0.01% 1 0.13% 0.5 0.20% AUCHENIPTERIDAE MUGILIDAE Parauchenipterus amblops 863 5.29% 112 15.09% 3.84 1.52% Mugil cf curema 2 0.01% 1 0.13% 0.45 0.18% PIMELODIDAE POLYNEMIDAE Rhamdia cf guatemalensis 509 3.12% 23 3.10% 5.15 2.03% Polydactylus 1 0.01% 0.00% 0.00% LORICARIIDAE P. approximans 1 0.01% 1 0.13% 0.4 0.16% Hypostomus panamensis 3 0.02% 1 0.13% 0.3 0.12% P. opercularis 109 0.67% 10 1.35% 4.2 1.66% cf ALBULIDAE unid.species 6 0.04% 1 0.13% 0.5 0.20% SCIAENIDAE 20 0.12% 0.00% 0.00% ALBULIDAE Bairdiella 1 0.01% 0.00% 0.00% Albula esuncula 52 0.32% 2 0.27% 1.1 0.43% B. ensifera 3 0.02% 2 0.27% 0.375 0.15% cf BATRACHOIDIDAE 1 0.01% 0.00% 0.00% Cynoscion 18 0.11% 0.00% 0.00% BATRACHOIDIDAE Cynoscion albus 17 0.10% 3 0.40% 5.35 2.11% Batrachoides pacifici 1 0.01% 1 0.13% 0.75 0.30% C. albus or stolzmanni 1 0.01% 1 0.13% 2.75 1.09% Daetor 1 0.01% 1 0.13% 0.2 0.08% C. stolzmanni 27 0.17% 4 0.54% 1.6 0.63% cf BELONIDAE unid species 1 0.01% 0.00% 0.00% C. cf stolzmanni 2 0.01% 2 0.27% 0.225 0.09% BELONIDAE Menticirrhus panamensis 1 0.01% 1 0.13% 0.55 0.22% Strongylura 1 0.01% 1 0.13% 0.25 0.10% Micropogonias altipinnis 22 0.13% 4 0.54% 4.15 1.64% Tylosurus 2 0.01% 2 0.27% 0.65 0.26% Ophioscion 1 0.01% 0.00% 0.00% PARALICHTHYIDAE Ophioscion scierus 7 0.04% 3 0.40% 0.55 0.22% Citharichthys gilberti 1 0.01% 1 0.13% 0.05 0.02% O. typicus 13 0.08% 3 0.40% 0.4 0.16% CARANGIDAE O. cf typicus 5 0.03% 0.00% 0.00% Alectes ciliaris 1 0.01% 1 0.13% 0.15 0.06% Paralonchurus dumerlii 1 0.01% 1 0.13% 0.8 0.32% Carangoides otrynter 10 0.06% 2 0.27% 0.85 0.34% Stellifer chrysoleuca 1 0.01% 1 0.13% 0.2 0.08% Caranx 4 0.02% 0.00% 0.00% SCOMBRIDAE C. caballus 39 0.24% 4 0.54% 0.7 0.28% Euthynnus lineatus 5 0.03% 1 0.13% 0.8 0.32% C. caninus 29 0.18% 6 0.81% 18.9 7.46% Scomberomorus sierra 7 0.04% 4 0.54% 1.95 0.77% Chloroscombrus orqueta 251 1.54% 13 1.75% 1.075 0.42% SPHYRAENIDAE

Table 8.1: Sitio Sierra: Fish species list (continued) 292

Estimated Estimated Body Genus and Species NISP % MNI % Body Mass % Genus and Species NISP % MNI % Mass % cf C. orqueta 2 0.01% 0.00% 0.00% Sphyraena ensis 2 0.01% 2 0.27% 0.95 0.37% Oligoplites 51 0.31% 0.00% 0.00% STERNARCHIDAE O. altus 9 0.06% 1 0.13% 1 0.39% Sternopygus dariensis 161 0.99% 9 1.21% 2.4 0.95% O. refulgens 25 0.15% 5 0.67% 1 0.39% STROMATEIDAE O. cf refulgens 1 0.01% 0.00% 0.00% Peprilus snyderi 2 0.01% 2 0.27% 0.35 0.14% O. saurus 1 0.01% 1 0.13% 0.25 0.10% SYNBRANCHIDAE Selar crumenophthalmus 25 0.15% 2 0.27% 0.35 0.14% Synbranchus marmoratus 117 0.72% 4 0.54% 0.725 0.29% Selene 354 2.17% 0.00% 0.00% TETRAODONTIDAE 6 0.04% 0.00% 0.00% Selene brevoortii 54 0.33% 4 0.54% 2.075 0.82% Sphoeroides 3 0.02% 0.00% 0.00% S. oersterdii 2 0.01% 1 0.13% 0.2 0.08% Sphoeroides annulatus 1 0.01% 1 0.13% 0.35 0.14% S. peruviana 1154 7.07% 50 6.74% 7.5 2.96% Guentheridia formosa 4 0.02% 2 0.27% 0.5 0.20%

Table 8.1: Sitio Sierra: Fish species list (continued)

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Results

Fish bones at Sitio Sierra have been exposed to differential heat as can be observed in Table

8.2. This suggests in general that 57.4% of the total sample of fish bones at Sitio Sierra were affected by fire.

NISP (not included %NISP Heating Category at Sitio Sierra freshwater taxa)

Completely calcined (White) 133 1.1

Partly calcined 87 0.7

Partly calcined/ partly carbonized 25 0.2

Carbonized (Black) 1178 9.6

Partly carbonized 9 0.07

Partly carmelized /partly carbonized 74 0.6

Carmelized (Red or Brown) 5262 42.9

Partly carmelized 101 0.8

Slightly carmelized 151 1.2

Unburnt 5221 42.6

Total 12241 100

Table 8.2: Differential heat exposure (NISP) and percentage of burnt fish remains from Sitio Sierra

294

295

The Clupeidae family (on charts 8.1a, 8. 1b and 8. 1c) shows that cranial skeleton bones outnumbered axial skeleton. The vertebral series indicates the precaudal vertebrae are missing. The relative element distribution of this family demonstrates that not all the regions are represented, the occipital and the otic regions are more frequent, and the branchial arch, pelvic girdle, vertebral column, branchial arch and the olfactory region the least frequent.

Figure 8.2: Survival index (SI) for Clupeidae fish remains from Sitio Sierra

This pattern is confirmed also by the survival index (Figure 8.2) which shows that both the otic and the occipital regions are overrepresented, showing larger values in comparison to other anatomical regions which values are smaller or underrepresented in

Sitio Sierra.

The Haemulidae pattern shows also that cranial bones are more abundant than the postcranial (Figure 8.3b). There are more thoracic vertebrae than the other types (Figure

8.3c). The relative element distribution (Figure 8.3a) suggests a larger representation of

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bones from the otic, the olfactory, pectoral girdle, occipital and vertebral regions which is corroborated by the survival index (Figure 8.4).

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The occipital, pectoral girdle, lateral, olfactory, and otic regions have each a survival index showing larger values in comparison to the other anatomical areas, which indicates that these regions are overrepresented (Figure 8.4).

Figure 8.4: Survival index (SI) for Haemulidae fish remains from Sitio Sierra

The family Carangidae describes a similar pattern. As shown in the graph 8.5b, the cranial skeleton outnumbered the postcranial bones. Almost all anatomical regions are represented at Sitio Sierra with the exception of the investing area, opercular, branchial arch, pelvic girdle and dorsal fin (Figure 8.5a). There is a predominance of thoracic vertebrae as the vertebral series chart (Figure 8.5c) indicates. 298

299

The situation that describe the survival index suggests that areas such as the caudal skeleton-vertebral column, occipital, anal fin, olfatory and lateral regions are over- represented in comparison to other anatomical regions. The investing, opercular, pelvic girdle, pectoral girdle and the branchial arch are the most underrepresented areas (Figure

8.6).

Figure 8.6: Survival index (SI) for Carangidae fish remains from Sitio Sierra

The family Ariidae describes a similar pattern. As shown in the graph 8.7b, the cranial skeleton outnumbered the postcranial bones. An examination of relative element representation of Ariidae reveals that areas such as the olfactory, occipital, lateral, hyoid arch, pectoral girdle, dorsal fin and vertebral column are represented in high numbers in comparison to the investing, otic, opercular, mandibular, pelvic region and the rest of the 300

axial skeleton (Figure 8.7a). There is a predominance of the vertebral complex as indicated in the vertebral series chart (Figure 8.7c).

The data from the Ariidae´s survival index (Figure 8.8) indicates that the dorsal fin, vertebral column, pectoral girdle, opercular, lateral, investing, and olfatory regions show larger values in comparison to the other regions. The underrepresentation of the caudal skeleton, branchial arch, the pelvic girdle, the pectoral fins and the opercular region is also remarkable.

Figure 8.8: Survival index (SI) for Ariidae fish remains from Sitio Sierra

The Sciaenidae family also repeats the pattern shown by other families. The graph

8.9b indicates that cranial bones are more frequent than postcranial ones. The vertebral series indicates the presence of thoracic vertebrae in comparison with the other types of vertebrae (Figure 8.9c).

. 302

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The relative element distribution suggests that bones from olfatory, lateral, caudal skeleton, and hyoid arch regions are more frequent than those that belong to the mandibular, investing, opercular , pelvic girdle and fins (Figure 8.9a). The survival index also indicates that bones from anal fin, pectoral girdle and pelvic girdle, branchial arch, mandicular, opercular, investing regions are underrepresented in compaarison with other regions such as olfatory, lateral, otic, occipital, hyoid arch, vertebral column and caudal skeleton, which are overrepresented at Sitio Sierra (Figure 8.10).

Figure 8.10: Survival index (SI) for Sciaenidae fish remains from Sitio Sierra

In the following Figures (Figures 8.11 to 8.13), I illustrate the vertebrae represented on Sitio Sierra sample. The top of each part of the pie chart is indicates the vertebrae 304

position. At the bottom is indicated the percentage. Families such as Ariidae, Clupeidae,

Haemulidae , and Sciaenidae, have adjacent caudal vertebrae (tail) and thoracic (head) vertebrae which are discarded as separate units after perhaps splitting the specimens. In some families such as Carangidae, vertebrae from diferent sections are thrown away.

Figure 8.11: Ariidae NISP vertebrae= 7 Carangidae NISP vertebrae=734

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Figure 8.12: Clupeidae NISP vertebrae= 74 Haemulidae NISP vertebrae=744

Figure 8.13: Sciaenidae=24

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AG-125´s Sample

The 622 fish skeletal elements recovered in this sample were identified to 22 families, 42 genera and 54 species (see Table 8.3 for details). In terms of NISP, the most abundant families in order of importance were Ariidae (37.14%), Clupeidae (13.99%), Carangidae

(12.38%), Tetraodontidae (10.13%), Haemulidae (9.65%), Sciaenidae (5.47%) and

Polynemidae (2.57%), in particular the genera Haemulopsis (NISP 18), Polydactylus(NISP

8), Ariopsis (NISP 8), Cathorops (NISP 23) and Cynoscion (NISP 15). Specifically the most abundant species were Ariopsis seemani (MNI 7), Opisthonema libertate (MNI 6), Selene peruviana

(5.2MNI 6), Haemulopsis leuciscus (MNI 6) and Chloroscombrus orqueta (MNI 6)(Cooke and

Jiménez 2004).

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Estimated Genus and Species NISP % MNI % Body Mass %

Elamosbranchs CARCHARHINIDAE 1 1 0.88% 600 3.10% C.limbatus 1 0 Rhizoprioniodon longurio 1 1 0.88% 1000 5.17% Total 3 0.48% DASYATIDAE Dasyatis longus 1 1 0.88% 400 2.07% Total 1 0.16% SPHYRNIDAE Sphyrna cf lewini 3 0 Total 3 0.48% UROLOPHIDAE Urotrygon cf asterias 1 1 0.88% Total 1 0.16% Teleosts ARIIDAE 192 Notarius 1 1 0.88% 130 0.67% Cathorops dasycephalus 2 2 1.77% 260 1.34% Ariopsis seemani 8 7 6.19% 1159 5.99% B. pinimaculatus 2 1 0.88% 1100 5.69% Cathorops 10 Cathorops cf fuerthii 1 C. multiradiatus 1 1 0.88% 130 0.67% C. cf multiradiatus 1 C. tuyra 4 3 2.65% 420 2.17% C. cf tuyra 4 3 2.65% 600 3.10% Sciades dowii 5 3 2.65% 480 2.48% Total 231 37.14% PARAUCHENIPTERIDAE Trachycorystes amblops 1 1 0.88% 5 0.03% Total 1 0.16% ALBULIDAE Albula nemomptera 1 1 0.88% 200 1.03%

Total 1 0.16% BELONIDAE 1 1 0.88% 40 0.21% Tylosurus 1 0 160 0.83% Total 2 0.32%

PARALICHTHYIDAE Citharichthys cf gilberti 17 4 3.54% 190 0.98% Total 17 2.73%

Table 8.3: Ag-125: Fish species list

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CARANGIDAE

C. caballus 6 3 2.65% 600 3.10% C. caninus 1 1 0.88% 400 2.07% Chloroscombrus orqueta 32 6 5.31% 230 1.19% O. altus 3 3 2.65% 230 1.19% Selene 1 S. peruviana 34 6 5.31% 790 4.08% Total 77 12.38% CENTROPOMIDAE Centropomus 1 C. armatus 1 0 C. cf robalito 1 1 0.88% 150 0.78% Total 3 0.48% CLUPEIDAE Opisthonema cf libertate 87 6 5.31% 770 3.98% Total 87 13.99% PRISTIGASTERIIDAE Opisthopterus dovii 6 3 2.65% 80 0.41% Total 6 0.96% GERREIDAE E. currani 1 1 0.88% 125 0.65% E. dowii 1 1 0.88% 130 0.67% Total 2 0.32% GOBIIDAE Gobiomorus maculatus 1 1 0.88% 35 0.18% Total 1 0.16% HAEMULIDAE 3 Orthopristis chalceus 38 4 3.54% 625 3.23% P. cf panamensis 1 1 0.88% 350 1.81% Haemulopsis 2 H. elongatus 2 1 0.88% 175 0.90% H. cf elongatus 2 1 0.88% 125 0.65% H. leuciscus 11 6 5.31% 1130 5.84% H. nitidus 1 1 0.88% 320 1.65% Total 60 9.65% OPHICTHIDAE E. brunneus 2 2 1.77% 600 3.10% Total 2 0.32% POLYNEMIDAE P. approximans 1 1 0.88% 300 1.55% P. opercularis 15 7 6.19% 1110 5.74% Total 16 2.57% SCIAENIDAE Bairdiella armata 1 1 0.88% 200 1.03%

Table 8.3: Ag-125: Fish species list (continued) 309

Cynoscion 5 Cynoscion albus 4 4 3.54% 1350 6.98% C. praedatorius 1 1 0.88% 175 0.90% C. stolzmanni 5 2 1.77% 275 1.42% Menticirrhus panamensis 7 3 2.65% 255 1.32% Micropogonias altipinnis 1 1 0.88% 150 0.78% Ophioscion scierus 1 1 0.88% 225 1.16% O. typicus 2 2 1.77% 250 1.29% O. cf typicus 1 L acclivis 1 1 0.88% 120 0.62% L. cf acclivis 1 1 0.88% 30 0.16% Stellifer oscitans 2 1 0.88% 175 0.90% Stellifer chrysoleuca 1 1 0.88% 100 0.52% Total 34 5.47% SCOMBRIDAE Scomberomorus sierra 1 1 0.88% 150 0.78% Total 1 0.16% SPHYRAENIDAE Sphyraena ensis 1 1 0.88% 150 0.78% Total 1 0.16% STROMATEIDAE Peprilus 2 Peprilus medius 2 1 0.88% 150 0.78% Peprilus snyderi 5 Total 9 1.45% TETRAODONTIDAE 2 Arothron 1 1 0.88% 120 0.62% Sphoeroides 2 Sphoeroides annulatus 2 1 0.88% 210 1.09% Guentheridia formosa 56 1 0.88% 110 0.57% Total 63 10.13% Grand total 100.00% 113 100.00% 19344 100.00%

Table 8.3: Ag-125: Fish species list (continued)

Results Fish bones at AG-125 have been exposed to differential heat as can be observed in Table

8.4. It suggests in general that 66% of the total sample of fish bones at AG-125 were affected by fire.

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Heating category at AG-125 NISP %

Completely calcined (White) 171 27.5

Partly calcined 12 1.9

Carbonized (Black) 152 24.4

Partly carmelized /partly carbonized 1 0.1

Carmelized (Red or Brown) 75 12.1

Unburnt 211 34

Total 622 100

Table 8.4: Differential heat exposure, (NISP) and percentage of burnt fish remains from Ag- 125

The Ag-125 pattern observed in the Ariidae family shows that cranial bones are less numerous than postcranial bones so this looks like a consumption site or the specimens were processed whole and gutted ventrally. Then the skull was kept intact to transport to a consumer site (Figure 8.14b). The vertebrae series indicates that the vertebral complex is more represented than the other types (Figure 8.14c). The relative element distribution of this family suggests that bones from the pectoral girdle, otic region, and vertebral column are more frequent than other anatomical regions. Investing, olfactory, mandibular, branchial arch and pelvic girdle´s bone are absent (Figure 8.14a)

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Figure 8.15: Survival index (SI) for Ariidae fish remains from Ag-125

This pattern is also confirmed by the survival index. The graph 8.15 shows that both the pectoral girdle , pectoral fin, vertebral colum, lateral and otic regions are over- represented (showing larger values). The branchial arch, mandicular, opercular, investing region and the pelvic gridle are poorly represented.

On the other hand, the Sciaenidae family also has more cranial than postcranial bones which suggest a production site (Figure 8.16b). Thoracic vertebrae are more frequent than the other type of vertebrae (Figure 8.16c). The element distribution´s chart at AG-125 313

shows that occipital, otic, and vertebral column bones are more frequent than the other anatomical regions (Figure 8.16a)

The survival index chart (Figure 8.17) suggests that the vertebral column, otic and the occipital anatomic regions (showing larger values) are over-represented in comparison to other anatomical regions which have smaller values.

Figure 8.17: Survival index (SI) for Sciaenidae fish remains from Ag-125

The Carangidae family at Ag-125 (on charts 8.18a, 8.18b and 8.18c) shows that cranial skeleton outnumbered postcranial one. The vertebral series indicates that there are more thoracic vertebrae than the other sectors.

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The relative element distribution of this family demonstrates that the caudal skeleton, anal and vertebral column are more frequent than the investing, branchial arch, pectoral girdle and pelvic girdle (Figure 8.18a). This pattern is confirmed also by the survival index which shows that the anal fin and vertebral column are overrepresented, showing values larger than other anatomical regions (Figure 8.19).

Figure 8.19: Survival index (SI) for Carangidae fish remains from Ag-125

The family Haemulidae describes a different pattern. As shown in the graph 8.20b, the cranial skeleton outnumbered axial bones. Not all anatomical regions are represented at Ag-

125.

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Occipital, otic, lateral region and vertebral column´s bones are more frequent than other anatomical regions (Figure 8.20a). There is a predominance of thoracic vertebrae as the vertebral series chart (Figure 8.20c) indicates.

Figure 8.18: Survival index (SI) for Haemulidae fish remains from Ag-125

The vertebral column, mandibular, otic and occipital regions show large values which indicate they are overrepresented in comparison to the other anatomical areas (Figure 8.21).

A higher proportion of Tetraodontidae bones were recorded at Ag-125(NISP 62) than at Sitio Sierra (NISP 6). At Ag-125 the sample of the Tetraodontidae family has more cranial than postcranial bones (Figure 8.22b). Thoracic vertebrae are more frequent than the other types of vertebrae (Figure 8.22c). The element distribution chart shows that lateral,

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and mandibular region bones are more frequent than the other anatomical regions (Figure

8.22a).

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Figure 8.23: Survival index (SI) for Tetraodontidae fish remains from Ag-125

The survival index chart (Figure 8.23) suggests that lateral and mandibular anatomic regions (showing larger values) are over-represented in comparison to other anatomical regions which have smaller values.

Finally, Clupeidae´s pattern shows that cranial bones slightly outnumbered the axial ones (Figure 8.24b). The vertebral series indicates a more frequent presence of precaudal vertebrae in comparison to caudal vertebrae and thoracic vertebrae (Figure 8.24c).

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Figure 8.25: Survival index (SI) for Clupeidae fish remains from Ag-125

The relative element distribution suggests occipital, otic, vertebral column and caudal skeleton are more frequent than other anatomical regions where their absence is outstanding

(Figure 8.23a). The survival index chart confirms that otic, occipital, caudal skeleton and vertebral column bones are over-represented in comparison to the absence of the other regions (Figure 8.24).

Because of the few vertebrae identified to a specific position, I only show the

Figures of the families Carangidae, Haemulidae and Sciaenidae (Figures 8.26 and 8.27). The top of each part of the pie chart the vertebrae position is indicated . At the bottom is indicated the percentage. These families, have adjacent caudal vertebrae (tail) and thoracic

(head) vertebrae which are discarded as separate units after perhaps filleting the specimens.

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Figure 8.26: Carangidae NISP vertebrae = 27 Haemulidae NISP vertebrae=13

Figure 8.27: Sciaenidae NISP vertebrae = 34

Butchering Units or Articulated Bones

Similar to the Vampiros sample, both Sitio Sierra and Ag-125 samples were studied in order to understand how fish were butchered (Lyman 1994; Reitz and Wing 1999). The following

Table(Table 8.5) illustrates the articulated bones in Sitio Sierra. There were no articulated 324

bones found at Ag-125. This information in conjuction with the element distribution provides some clues about how people butchered and used fish.

Family Bones Region NISP

Ariidae Articular-Dentary Lateral 2

Ceratohyal-Epihyal Hyoid arch 8

Ceratohyal-Epihyal-Hypohyal Hyoid arch 3

Ceratohyal-Hypohyal Hyoid arch 12

Epiotic/Pteriotic/Scale bone/Frontal Otic-Investing 4

Frontal/Supracleithrum/Posttemporal/Supraoccipi Otic-Investing 4 tal

Pteriotic-Epiotic Otic 2

Sphenotic-Alisphenoid Otic 2

Supraoccipital-Pteriotic Otic 2

Vomer-Frontal Investing-Olfatory 12

Carangidae Ceratohyal-Epihyal Hyoid arch 2

Clupeidae Prootic-Pteriotic Otic 2

Caudal vertebrae 253

Centropomidae Frontal Investing 2

Haemulidae Supraoccipital-Epiotic Occipital-Otic 10

Epiotic- Exocipital-Prootic-Pteriotic Otic 8

Epiotic- Exocipital-Prootic-Pteriotic-Sphenotic Otic 5

Neurocranium 28

Polynemidae Ceratohyal-Epihyal Hyoid arch 2

Sphenotic-Prootic Otic 2 Table 8.5: Sitio Sierra: articulated bones by anatomic region

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Articulated fish bones found at Sitio Sierra are represented in Ariidae, Carangidae,

Clupeidae, Centropomidae, Haemulidae Polynemidae. These include a complete neurocranium and several units on the skull, which suggest that the head was removed and discarded as a whole (occipital, investing, occipital and otic regions) or adjacent regions

(hyoid arch and lateral regions). As shown in Table 8.5, most of articulated vertebrae were identified to the Clupeidae family corresponding to caudal vertebrae.

Cut marks

As observed on the Vampiros samples and table 7.22 following Belcher´s (1998) Fisher

(1995), Seetah (2006), and Zohar´s (2003) criteria, I observed each bone of Sitio Sierra and

Ag-125 to identify cut marks, recording their location, number and morphology.

In general both sites have few cut marks in comparison to Vampiros rockshelters

(Figure 8.28 and Table 8.6) and similar to Vampiros the cut marks tended to be shallow and small striations (cut) caused by knife, impact is like a percussion pit with a circular perforation or depresion for using a hammerstone or perhaps a polished axe , crushing is a splintered marging of the bone from a hammerstone or perhaps a polished axe, and chopping mark is smooth entry and fractured exit points for using a hammerstone or perhaps a polished axe.

In Sitio Sierra´s Ariidae sample, cut marks and fractures are concentrated on ceratohyal, cleithrum, pectoral spines, hyomandibular, vertebrae and ethmoid on a lateral side. They

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exhibited three patterns: First, fine transverse striations produced by a knife (ceratohyal, hypohyal, vertebrae _Figure 8.28a-, epyhial, supracleithrum). According to Willis et al

(2008:1441) cut marks of this type on the supracleithrum result from butchering activities when people decapitate catfish. The marks on ceratohyal, epihyal and hypohyal are result from eviscerating fish. Second, smooth entry and fractured exit points (Figure 8.28b,c) or splintererd marging of the bone for splitting (cleithrum, ethmoid, and hypohyal,). Third there are circular perforations or depresions for using a hammerstone or perhaps a polished axe (Figure 8.28b). Finally, marks or fractures on pectoral and dorsal spines with splintered marging of the bone, produced by cracking the spines to avoid injuries.

Figure 8.28: a. cut mark on Ariidae´s caudal vertebrae, b. Ariidae´s cleithrum showing chopped and impact marks and, c. Sciaenidae´s premaxilla chopped.

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Site Family Bone # Portion Morphology

Sitio Ariidae Hypohyal- 1 Anterior-Dorsal Incision Sierra Ceratohyal

Ceratohyal 1 Lateral Crushed. Irregular Cleithrum 4 Lateral-Mid Chopped- Straight-Impact Ethmoid 1 Lateral Crushed- Irregular Pectoral 1 Posterior Crushed - spine Irregular Hyomandibu 1 Lateral Chopped- lar Straight Dorsal spine 1 Posterior Crushed- irregular Vertebra 1 Mid Incision Hypohyal 1 Anterior Crushed- Irregular Sitio Carangidae Articular 3 Anterior Crushed- Sierra Irregular Ceratohyal 1 Posterior Incision Epihyal 1 Lateral Incision Supracleithru 1 Lateral Incision m Sitio Haemulidae Cleithrum 1 Mid Chopped- Sierra Straight Parasphenoid Mid Chopped- Straight Ceratohyal 1 Lateral Chopped- Straight Anal spine 2 Anterior Crushed- Irregular Sitio Sciaenidae Premaxilla 2 Anterior-lateral Chopped- Sierra Straight Dentary 1 Anterior-lateral Chopped- Straight Pterygiophor 1 Anterior Chopped- e anal Straight Ag- Sciaenidae Anal spine 1 Anterior Crushed- 125 Irregular Ag- Tetraodontidae Dentary 1 Lateral-Mid Incision 125

Table 8.6: Distribution of cut marks at Sitio Sierra and Ag-125

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Sitio Sierra´s Carangidae samples present transverse cut marks localized especially in the ceratohyal, epyhial, supracleithrum, and articular on the lateral side. They describe two patterns: first; fine striations produced by a knife (supracleithrum, ceratohyal, epihyal) for removing the head and gutting and second; splintered marging of the bone from a hammerstone or perhaps a polished axe on the angular.

Few cut marks were observed in Haemulidae bones. These were found mainly in the cleithrum, parasphenoid, ceratohyal and anal spine. These are transversal cuts or fractures related with decapitation and evisceration (smooth entry and fractured exit points on the cleithrum and ceratohyal) or cracking anal spines (splintered marging of the bone) to avoid injuries.

Sciaenidae bones present also evidence of cut marks. These are transverse cuts located on the anterior portion of each bone. They were found mainly in premaxilla, dentary and Pterygiophore anal. According to Belcher (1998:179) the marks on a lateral side on cranial elements are common on fresh fish consumption when fish is segmented into similar size pieces.

Damage

The first step of the analysis was to differentiate beween ancient damage or fractures from recent ones (Table 7.20). A bone with recent damage has sharp edges, and a clean, white appearance or light color that contrasts with the darker color of the bone's cortical surface. Bones with ancient damage lack of these characteristics.

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Later, I observed fragmentation patterns from Sitio Sierra and AG- 125 selecting the main families Ariidae, Carangidae, Clupeidae, Haemulidae, Sciaenidae and Tetraodontidae. I grouped the samples according to element.

NISP % Complete 3352 27.38% Predepositional 4922 40.21% Postdepositional 3956 32.32% N/A 11 0.09% Total 12241 100.00%

Table 8.7: Frequency (NISP) and percentage of predepositional and postdepositional fractures and completeness at Sitio Sierra

I examined the relative frequency of damaged bone at Sitio Sierra (Table 8.7). Most of the sample is fragmented (NISP 8889, 72.62%) either for pre-depositional (i.e. cooking, butchering) or post-depositional events (i.e. burning, trampling). Six vertebrae present evidence of trampling showing a longitudinal deformation.

Sitio Sierra 1/8" NISP % 100% 3352 26.94% 5% 0 0.00% 25% 1039 8.35% 40% 1214 9.76% 50% 962 7.73% 75% 4320 34.72% 90% 1545 12.42% Indet. 9 0.07% Total 12441 100.00%

Table 8.8: Frequency (NISP) and percentage of fragmentation at Sitio Sierra

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In Sitio Sierra´s sample the bones were complete or almost complete (between size categories 100% to 75% ca. 26.94%, 34.72% and 12.42% respectively, see Table 8.8).

However highly fragmented bones (between size 5% and 45%) were also identified.

Fractures

Considering the bones differential state of fragmentation, I address the bone's state of fragmentation for Sitio Sierra´s sample. The weighted mean index of fragmentation

(WMI) was examined according to the most abundant families: Ariidae, Carangidae,

Haemulidae, Clupeidae and Sciaenidae.

Anatomic region 100% 5% 25% 40% 50% 75% 90% WMI Olfactory 7.17 0.00 3.25 5.50 4.60 1.00 5.60 18.27 Occipital 9.00 0.00 7.00 6.00 11.50 1.50 3.50 23.18 Otic 23.80 0.00 3.20 6.40 7.75 1.00 4.80 36.11 Investing 17.00 0.00 2.00 22.00 10.00 2.00 10.00 41.80 Lateral 11.75 0.00 4.00 8.67 4.50 1.00 5.25 23.94 Opercular 9.50 0.00 6.33 3.33 2.33 1.00 3.00 17.03 Mandibular 8.00 0.00 0.00 2.50 2.50 1.50 3.00 14.08 Hyoid Arch 12.00 0.00 4.00 4.25 3.50 1.00 1.33 18.40 Branchial Arch 6.71 0.00 1.00 1.00 2.00 1.00 4.80 13.43 Pectoral Girdle 7.50 0.00 22.00 25.75 15.00 5.33 3.00 37.50 Pelvic Girdle 1.00 0.00 3.00 6.00 0.00 1.00 0.00 4.90 Vertebral Column 123.50 0.00 77.00 22.50 2.50 4.00 4.00 159.60 Caudal skeleton 150.50 0.00 2.00 4.00 6.00 11.00 0.00 163.85 Pectoral fin 24.00 0.00 21.50 15.50 9.00 5.00 7.00 50.13 Dorsal fin 1.50 0.00 5.00 8.00 5.00 1.00 0.00 9.20

Table 8.9: Frequency (%) of Ariidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Sitio Sierra

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In the Ariidae family most of the anatomic areas are highly fragmented (<40%) with the exception of the vertebral column (159.60%) and caudal skeleton (163.85%) which indicate their best preservation (Table 8.9).

Anatomic region 100% 5% 25% 50% 60% 75% 90% WMI Olfactory 12.00 0.00 0.00 4.67 4.33 4.00 5.50 23.98 Occipital 5.00 0.00 15.00 5.50 3.50 1.00 4.00 17.05 Otic 8.50 0.00 0.00 0.00 1.00 0.00 3.67 12.30 Investing 0.00 0.00 3.00 5.00 0.00 0.00 0.00 2.75 Lateral 1.00 0.00 8.00 6.40 2.33 2.00 1.50 9.58 Opercular 0.00 0.00 0.00 1.00 0.00 0.00 0.00 0.40 Mandibular 2.00 0.00 3.50 5.33 3.00 1.00 2.67 9.66 Hyoid Arch 4.25 0.00 1.50 11.00 5.00 1.00 4.00 15.88 Branchial Arch 2.00 0.00 1.00 1.00 1.50 0.00 0.00 3.40 Pectoral Girdle 2.67 0.00 9.00 8.00 3.00 0.00 2.50 11.87 Pelvic Girdle 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Vertebral Column 38.30 0.00 0.00 7.30 19.50 31.00 135.50 196.17 Caudal skeleton 40.00 0.00 11.00 2.00 7.50 51.00 66.00 144.95 Anal fin 0.00 0.00 7.00 26.00 8.00 0.00 41.00 53.05 Pectoral fin 0.00 0.00 0.00 0.00 2.00 0.00 2.00 2.80

Table 8.10: Frequency (%) of Carangidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Sitio Sierra.

Similar to the previous family, for the Carangidae family the WMI values show that most of the anatomic regions are highly fragmented (<40%); however, the vertebral column and caudal skeleton are the best preserved (> 50% see Table 8.10).

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Anatomic region 100% 5% 25% 50% 60% 75% 90% WMI Olfactory 0.00 0.00 0.00 0.00 0.00 0.00 3.00 2.70 Occipital 49.50 0.00 0.00 0.00 0.00 0.00 0.00 49.50 Otic 64.83 0.00 2.00 2.00 1.50 0.00 1.00 67.78 Investing 3.00 0.00 27.00 28.00 16.00 0.00 2.00 30.75 Lateral 4.00 0.00 0.00 7.00 1.00 3.00 0.00 9.55 Opercular 0.00 0.00 7.00 16.00 3.00 0.00 0.00 9.65 Mandibular 0.00 0.00 1.00 0.00 1.00 0.00 3.00 3.45 Hyoid Arch 2.00 0.00 11.00 27.00 9.50 3.00 1.33 23.75 Branchial Arch 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Pectoral Girdle 13.00 0.00 11.00 12.00 4.00 2.00 2.00 25.85 Pelvic Girdle 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Vertebral Column 9.50 0.00 0.00 0.00 0.00 1.00 0.00 10.25 Caudal skeleton 107.67 0.00 0.00 0.00 0.00 108.00 2.00 190.47

Table 8.11: Frequency (%) of Clupeidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Sitio Sierra.

In the Clupeidae family, the branchial arch and the pelvic girdle are absent in the sample, which explains the low value of WMI (0%), and therefore, their underrepresentation is not cause for fragmentation. In general, most of the anatomic regions are highly fragmented (<40%); however, the otic region and caudal skeleton are the best preserved (>

50% see Table 8.11).

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Anatomic region 100% 5% 25% 50% 60% 75% 90% WMI Olfactory 13.00 0.00 7.50 14.67 8.25 5.67 5.50 34.07 Occipital 22.33 0.00 6.00 16.33 8.67 12.00 7.33 50.30 Otic 44.80 0.00 9.00 1.00 9.00 26.00 21.33 90.65 Investing 9.50 0.00 20.00 23.00 13.00 1.00 5.00 35.45 Lateral 2.20 0.00 9.20 19.60 3.20 3.00 3.00 18.89 Opercular 5.00 0.00 45.00 4.00 4.33 5.00 1.67 25.27 Mandibular 13.50 0.00 6.00 6.50 11.00 4.00 2.00 27.90 Hyoid Arch 16.00 0.00 14.00 23.50 24.00 7.00 3.00 48.85 Branchial Arch 3.50 0.00 0.00 2.00 6.50 5.00 1.67 12.80 Pectoral Girdle 6.67 0.00 11.00 27.00 12.33 9.67 12.00 44.43 Pelvic Girdle 0.00 0.00 22.00 0.00 0.00 0.00 0.00 5.50 Vertebral Column 83.33 0.00 0.00 0.00 3.00 106.67 58.00 217.03 Caudal skeleton 18.50 0.00 0.00 0.00 2.00 23.00 258.00 268.95 Anal fin 17.00 0.00 10.00 11.00 10.50 1.00 10.00 38.90 Dorsal fin 2.00 0.00 0.00 1.00 0.00 0.00 3.00 5.10 Pectoral fin 0.00 0.00 0.00 0.00 0.00 1.00 9.00 8.85

Table 8.12: Frequency (%) of Haemulidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Sitio Sierra.

The Haemulidae fracture pattern shows that most of the anatomic region is highly fragmented (<40%); however, the otic region, vertebral column and caudal skeleton are the best preserved (> 50% see Table 8.12).

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Anatomic region 100% 5% 25% 40% 50% 75% 90% WMI Olfactory 2.00 0.00 2.00 1.00 0.00 0.00 1.00 3.80 Occipital 1.00 0.00 1.00 1.00 1.00 0.00 1.00 3.05 Otic 1.75 0.00 0.00 0.00 0.00 0.00 0.00 1.75 Investing 1.00 0.00 0.00 0.00 0.00 1.00 1.00 2.65 Lateral 0.00 0.00 1.25 1.67 1.67 0.00 1.67 3.31 Opercular 0.00 0.00 1.00 0.00 0.00 0.00 3.00 2.95 Mandibular 1.00 0.00 0.00 1.00 0.00 0.00 0.00 1.40 Hyoid Arch 1.75 0.00 4.00 0.00 0.00 0.00 1.00 3.65 Branchial Arch 1.00 0.00 0.00 0.00 0.00 0.00 0.00 1.00 Pectoral Girdle 1.33 0.00 0.00 1.00 1.00 0.00 1.00 3.13 Pelvic Girdle 0.00 0.00 0.00 1.00 1.00 0.00 0.00 0.90 Vertebral column 4.00 0.00 0.00 0.50 0.50 2.50 0.50 6.78 Caudal skeleton 8.33 0.00 0.00 0.00 0.00 5.00 0.00 12.08 Pectoral fin 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Anal fin 0.00 0.00 1.00 1.00 0.00 1.00 0.00 1.40

Table 8.13: Frequency (%) of Sciaenidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Sitio Sierra.

Similar to previous families, for the Sciaenidae family the WMI values show that most of the anatomic region are highly fragmented (<40%), however the caudal skeleton is the best preserved anatomical region (Table 8.13).

For comparison between the results, I ranked the worst preserved skeletal elements

(<40 %) with NISP larger than 10, in each taxonomic group. While most of the best preserved material in Sitio Sierra corresponds to vertebrae, their common type of fracture is related to the loss of haemal and neural spines (Table 8.14).

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Proximal Proximal Proximal Distal Distal Almost Lateral Bone WMI Rank end end/medial medial Medial Medial end/medial complete Caudal Lateral Centrum centrum BAH 9.50 3 2 2 3 BAM 5.50 1 9 1 DE 11.00 4 12 2 1 7 EXO 11.75 5 2 Ariidae PRF 8.75 2 2 3 3

ANG 10.15 5 2 23 DEN 6.70 2 13 5 EPH 10.80 8 1 4 EPI 12.00 10 MAX 6.15 1 4 2 4 1 1 1 PAL 7.40 3 7 4 2 PM 11.65 9 14 12 PRT 9.80 4 2 PTE 10.30 7 1 3 Carangidae QUA 10.20 6 7 9 1 3

CER 11.15 1 12 1 6 1 OPE 11.55 2 26 SPH 13.10 5 1 2 PENULTIMATE 12.40 4 1 2 Clupeidae THORACIC 12.75 3 1

BAM 5.5 1 22 CEB 10.95 6 2 8 2 2 1 DEN 6.85 2 5 10 EP 8.85 4 1 9 ETH 8.9 5 1 7 INT 12.7 7 11 1 Haemulidae URO 7.2 3 3 5 3

Sciaenidae PRECAUDAL 10.25 1 95 173 6

Table 8.14: Weighted mean index (WMI) of highly fragmented bones and portion (NISP) for Sitio Sierra sample

Highly fragmented bones (<40%) are basihyal, basipterygium, dorsal spine, exoccipital, and prefrontal for Ariidae; angular, dentary, epihyal, epiotic, maxilla, palatine, premaxilla, prootic, pterotic, and quadrate for Carangidae. The Clupeidae´s ceratohyal, opercle, sphenotic, penultimate and thoracic vertebrae are highly fragmented. For the

Haemulidae family the following bones are highly fragmented: basipterygium, ceratobranchial, dentary, pectoral spine, ethmoid, interopercle and urohyal; and finally the precaudal vertebrae for Sciaenidae are fragmented. These bones (see Table 8.14) show several fractured portions due to perhaps transverse cuts more than Vampiros. It seems based on this pattern of fractures and cut marks (lateral side) mainly on cranial elements that fish at Sitio Sierra was chopped into several portions for cooking (Belcher 1998: 179). 336

Typical fractures observed for all families are on the cleithrum. This is due to a transverse cut either to facilitate salt absorption or to remove the head (Belcher 1998; Zohar and Cooke 1997)(Table 8.15).

Grand Zone Ariidae Carangidae Clupeidae Haemulidae Sciaenidae Total

proximal end 39 3 40 1 83

proximal end/medial 6 4 8 18

proximal medial 2 2 6 10

Medial 49 16 2 29 1 97

distal/medial 18 7 6 31

distal end/medial 3 1 2 15 21

Complete 1 1

Lateral 6 6

Grand Total 124 30 7 104 2 267

Table 8.15: Types of fractures observed on the cleithrum on the most abundant families at Sitio Sierra

The fragmentation patterns for AG- 125 indicate that almost 265 (42.60%) bones are fragmented by post-depositional events (i.e. burning, storing) and 207 (33.28%) are 337

fragmented by pre-depositional events (i.e. butchering, cooking). Only 150 (24.12%) bones are complete (see Table 8.16). There was no evidence of trampling; none of the vertebrae present longitudinal deformation.

Complete Predepositional Postdepositional Total NISP 150 207 265 622 % 24.12% 33.28% 42.60% 100.00%

Table 8.16: Frequency (NISP) and percentage of predepositional and postdepositional fractures and completeness at Ag-125

Although Ag-125´s sample is small, the bones were complete or almost complete ( between size categories 100% to 75% ca. 24.12%, 13.02% and 26.53% respectively, see

Table 8.17). However, highly fragmented bones (between size 5% and 40%) were also identified.

Size class 100% 5% 25% 50% 60% 75% 90% Grand Total NISP 150 0 121 47 58 165 81 622

% 24.12% 0.00% 19.45% 7.56% 9.32% 26.53% 13.02% 100.00%

Table 8.17: Frequency (NISP) and percentage of fragmentation at Ag-125

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Considering the bones‘ differential state of fragmentation, I will address the bones‘ state of fragmentation for Ag-125´s sample. The weighted mean index of fragmentation

(WMI) will be examined according to the most abundant families: Ariidae, Carangidae,

Haemulidae, Clupeidae, Sciaenidae and Tetraodontidae.

Anatomic region 100% 5% 25% 50% 60% 75% 90% WMI Olfactory 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Occipital 0.00 0.00 0.00 0.00 0.00 0.00 1.00 0.90 Otic 7.00 0.00 0.00 1.00 2.00 0.00 1.00 9.30 Investing 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Lateral 2.00 0.00 2.00 2.00 0.00 0.00 0.00 3.30 Opercular 0.00 0.00 0.00 1.00 0.00 0.00 0.00 0.40 Mandibular 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Hyoid Arch 1.00 0.00 1.00 1.00 0.00 1.00 0.00 2.40 Branchial Arch 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Pectoral Girdle 0.00 0.00 5.67 1.00 1.00 1.00 0.00 3.07

Pelvic Girdle 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Vertebral Column 6.00 0.00 4.00 0.00 1.00 8.00 1.00 14.40 Caudal skeleton 36.00 0.00 0.00 0.00 0.00 14.00 5.00 51.00 Pectoral fin 0.00 0.00 5.00 1.00 0.00 3.00 0.00 3.90 Dorsal fin 1.00 0.00 2.00 1.00 1.00 0.00 0.00 2.40

Table 8.18: Frequency (%) of Ariidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Ag-125

In the Ariidae family, the olfatory, investing, mandibular, branchial arch and the pelvic girdle are absent regions in the sample, which explains the low value of WMI (0%), and therefore, their underrepresentation is not caused by fragmentation. In general, most of the anatomic regions are highly fragmented (<40%), however the caudal skeleton is the best preserved (> 50% see Table 8.18).

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Anatomic region 100% 5% 25% 50% 60% 75% 90% WMI Olfactory 2.00 0.00 0.00 0.00 0.00 0.00 1.00 2.90 Occipital 1.00 0.00 0.00 0.00 0.00 0.00 1.00 1.90 Otic 1.00 0.00 0.00 0.00 0.00 0.00 0.00 1.00 Investing 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Lateral 0.00 0.00 1.00 1.00 1.00 0.00 0.00 1.15 Opercular 0.00 0.00 1.00 0.00 0.00 0.00 0.00 0.25 Mandibular 0.00 0.00 1.00 0.00 0.00 0.00 0.00 0.25 Hyoid Arch 0.00 0.00 1.00 0.00 0.00 0.00 1.00 1.15 Branchial Arch 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Pectoral Girdle 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00

Pelvic Girdle 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Vertebral Column 3.00 0.00 0.50 0.00 1.00 3.00 2.50 8.13 Caudal skeleton 1.50 0.00 0.00 0.00 2.00 2.00 3.00 6.70 Anal fin 1.00 0.00 2.00 1.00 0.00 0.00 0.00 1.90

Table 8.19: Frequency (%) of Carangidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Ag-125

The Carangidae family the WMI values show that most of the anatomic regions are highly fragmented (<40%). The investing, pectoral girdle, branchial arch and the pelvic girdle are absent regions in the sample, which explains the low value of WMI (0%) (see

Table 8.19).

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Anatomic region 100% 5% 25% 50% 60% 75% 90% WMI Olfactory 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Occipital 1.00 0.00 0.00 0.00 0.00 0.00 0.00 1.00 Otic 2.25 0.00 0.00 0.00 0.00 0.00 0.00 2.25 Investing 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Lateral 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Opercular 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Mandibular 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Hyoid Arch 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Branchial Arch 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Pectoral Girdle 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00

Pelvic Girdle 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Vertebral Column 4.00 0.00 0.00 0.00 0.00 3.00 0.00 6.25 Caudal skeleton 0.00 0.00 0.00 0.00 0.00 25.00 28.00 43.95

Table 8.20: Frequency (%) of Clupeidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Ag-125

For the Clupeidae family, the olfactory, investing, lateral, opercular, mandibular, hyoid arch, pectoral girdle, branchial arch and the pelvic girdle are absent regions in the sample, which explains the low value of WMI (0%) and therefore, their underrepresentation. In general, most of the anatomic regions are highly fragmented (<40%), with the exception of the caudal skeleton, which is the best preserved (43.95% see Table 8.20).

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Anatomic region 100% 5% 25% 50% 60% 75% 90% WMI Olfactory 0.00 0.00 0.00 1.00 0.00 0.00 0.00 0.40 Occipital 1.00 0.00 0.00 0.00 0.00 0.00 2.00 2.80 Otic 2.50 0.00 0.00 0.00 1.00 0.00 1.00 3.90 Investing 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Lateral 0.00 0.00 1.00 0.00 2.00 0.00 0.00 1.25 Opercular 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Mandibular 1.00 0.00 0.00 0.00 1.00 0.00 1.00 2.40 Hyoid Arch 0.00 0.00 0.00 0.00 0.00 1.00 1.00 1.65 Branchial Arch 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Pectoral Girdle 0.00 0.00 0.00 0.00 0.00 0.00 1.00 0.90

Pelvic Girdle 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Vertebral Column 2.67 0.00 0.00 0.00 0.00 2.00 1.00 5.07 Caudal skeleton 3.00 0.00 0.00 0.00 0.00 7.00 2.00 10.05 Anal fin 0.00 0.00 1.00 0.00 2.00 1.00 0.00 2.00

Table 8.21: Frequency (%) of Haemulidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Ag-125

The few bones belonging to the Haemulidae family show that most of the anatomic regions are highly fragmented (<40%) or with low WMI´s values (0%) such as the investing, opercular, branchial arch and pelvic girdle and therefore their underrepresentation (Table

8.21).

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Anatomic region 100% 5% 25% 40% 50% 75% 90% WMI Olfactory 0.00 0.00 0.00 0.00 0.00 0.00 0.00 5.30 Occipital 1.00 0.00 0.00 0.00 0.00 0.00 1.00 5.62 Otic 3.00 0.00 0.00 0.00 0.00 0.00 1.00 2.50 Investing 0.00 0.00 0.00 0.00 0.00 0.00 0.00 1.25 Lateral 0.00 0.00 1.00 0.00 3.00 0.00 0.00 5.24 Opercular 0.00 0.00 0.00 0.00 0.00 0.00 0.00 2.15 Mandibular 0.00 0.00 0.00 0.00 0.00 0.00 0.00 5.05 Hyoid Arch 0.00 0.00 0.00 0.00 0.00 1.00 0.00 7.20 Branchial Arch 0.00 0.00 0.00 0.00 0.00 0.00 0.00 5.65 Pectoral Girdle 0.00 0.00 0.00 0.00 0.00 0.00 0.00 9.10 Pelvic Girdle 0.00 0.00 0.00 0.00 0.00 0.00 0.00 1.75 Vertebral column 3.00 0.00 0.00 0.00 0.00 0.00 1.00 12.05 Caudal skeleton 1.00 0.00 0.00 0.00 1.00 4.00 2.00 45.65 Pectoral fin 0.00 0.00 0.00 0.00 0.00 0.00 0.00 2.15 Anal fin 0.00 0.00 0.00 0.00 0.00 1.00 0.00 12.00

Table 8.22: Frequency (%) of Sciaenidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Ag-125

For Sciaenidae family, the WMI values show that most of the anatomic regions are highly fragmented (<40%); however, the caudal skeleton (45.65%) is the best preserved anatomical region (Figure 8.22).

Anatomic region 100% 5% 25% 40% 50% 75% 90% WMI Olfactory 0.00 0.00 0.00 0.00 0.00 0.00 0.00 5.12 Occipital 0.00 0.00 0.00 0.00 0.00 0.00 1.00 2.57 Otic 1.00 0.00 0.00 0.00 0.00 0.00 0.00 5.50 Investing 0.00 0.00 0.00 0.00 0.00 0.00 0.00 12.20 Lateral 1.33 0.00 1.00 1.00 1.67 2.50 1.00 13.08 Opercular 0.00 0.00 2.00 1.00 0.00 0.00 0.00 10.78 Mandibular 3.00 0.00 3.00 2.00 0.00 0.00 0.00 12.30 Hyoid Arch 0.00 0.00 0.00 0.00 0.00 0.00 1.00 16.85 Branchial Arch 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Pectoral Girdle 0.00 0.00 0.00 0.00 0.00 0.00 0.00 10.68 Vertebral column 1.50 0.00 0.00 0.00 0.00 2.00 0.00 22.94 Caudal skeleton 0.00 0.00 0.00 0.00 0.00 0.00 0.00 18.30

Table 8.23: Frequency (%) of Tetraodontidae skeletal elements by structure, according to the seven fragmentation categories and their weighted mean index (WMI), in Ag-125

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Finally, the Tetraodontidae family (Table 8.23) shows that most of the anatomic regions

are highly fragmented (<40%) and the branchial arch presents a low WMI´s values (0%

Proximal Distal Almost Anterior Lateral Bone WMI Rank end Medial end/medial complete Centrum centrum centrum CLE 3.25 1 4 8 Ariidae UND.SPINE 12.75 2 2 29 6

Carangidae VERT FRAG 11.55 1 8 6

Clupeidae VERT FRAG 3.30 1 3 8

Haemulidae CAUDAL 10.05 1 2 7

Sciaenidae VERT FRAG 9.35 1 11 3 2

Tetraodontidae VERT FRAG 10.95 1 25 4 2

Table 8.24: Weighted mean index (WMI) of highly fragmented bones and portion (NISP) for Ag-125 sample

Similar to Sitio Sierra sample, I ranked the worst preserved skeletal elements (<40 %) with

NISP larger than 10 in each taxonomic group in Ag-125. The most frequently damaged bones in AG-125 (see Table 8.24) are the vertebrae for Carangidae, Clupeidae, Haemulidae,

Sciaenidae and Tetraodontidae. The common type of fracture is related to the lost of haemal and neural spines. Wheeler and Jones (1988:108) consider this break off as a result of post- depositionally trampling or other taphonomic agents. I did not consider this fracture as postdepositional since I did not observe a light coloration.

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Typical fractures observed for Ariidae are on the cleithrum and spines. Cleithra fractures are due to a transverse cut either to facilitate salt absorption or to remove the head.

Summary

Both Sitio Sierra and Ag- 125 have been nice locations for settlement because of the multiple natural resources for fishing near the estuary. These sites show that most of the marine taxa consisted of three species which often occur in large schools: pacific thread hearing (cf Opisthonema libertate), brassy grunt (Orthopristis chalceus) and pacific moonfish (Selene peruviana)(Cooke, et al. 2008; Weiland 1984).

To understand how Pre-Columbian inhabitants processed and/or discarded fish in both sites, the abundance of elements (%NISP) was organized in a graphic manner, each color indicates a percentage being less than 1% (red) to >5% (purple). A black mark indicates fractures and a purple star indicates cut marks.

At Sitio Sierra, the pattern observed for the Ariidae family indicates that people were discarding more cranial bones than axial ones, especially those from the olfactory, pectoral girdle and caudal skeleton. The few anterior/posterior/ lateral cut marks indicated that people were removing the flesh (ceratohyal, epihyal, hipohyal) and breaking dorsal spines

(Figure 8.29).

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Figure 8.29: Ariidae specimens from Sitio Sierra. NISP= 2535

There is a low frequency of first dorsal spines and fin bones. Zones such as the otic area including otoliths and the branchial archs are underrepresented. This might be as result of butchering practices. Those less frequent bones were discarded somewhere else perhaps at a processing site. Belcher (1998) observed similar patterns in dried fish processing samples.

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Figure 8.30: Carangidae specimens from Sitio Sierra. NISP= 2229

People who lived at Sitio Sierra were disposing more Carangidae cranial than axial bones (Figure 8.30). In fact, most of the bones from skull, pectoral girdle, fins and the mid- section (pre-caudal vertebrae) are missing and the sample comprises mainly thoracic vertebrae. Those missing bones perhaps were discarded somewhere else such as a processing site. The few cut marks are located on branchial arch as a result of removing or chopping the head

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In the case of the Clupeidae family, all anatomical areas are underrepresented at Sitio Sierra with the exception of the occipital and otic (orange) regions. The missing regions were discarded somewhere else or ate (Figure 8.31).

Figure 8.31: Clupeidae specimens from Sitio Sierra. NISP= 3824

The Sciaenidae pattern observed indicates that some anatomical regions are missing such as the fins and the branchial arch (Figure 8.32). People gutted and removed fins and deposited them somewhere else like a processing site. The few cut marks are located on the lateral area, and their posterior location might suggest other chopping acts different to those for dried fish (Belcher 1998).

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Figure 8.32: Sciaenidae specimens from Sitio Sierra. NISP= 158

For the Haemulidae family (Figure 8.33), the pattern suggests that Pre-Columbian inhabitants were discarding all anatomical regions, especially the investing region and thoracic vertebrae. However, the few cut-marks found were located on the branchial arch.

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Figure 8.33: Haemulidae specimens from Sitio Sierra. NISP= 3619

The site AG-125 has a small sample which did not permit a definitive statement; however, the Ariidae sample (Figure 8.34) suggests that bones from investing, olfactory, mandibular, branchial arch and pelvic girdle regions are absent, and cut marks were not observed. Perhaps those bones were tossed somewhere else. Most of the sample comprises bones from fins, otoliths and the pectoral girdle, which implies that the Pre-Columbian inhabitants were gutting the fish and discarding fins at this site.

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Figure 8.34: Ariidae specimens from Ag-125. NISP= 232

The Carangidae pattern at AG-125 indicates that there are more axial than cranial bones. Anatomical regions such as investing, branchial arch, pectoral girdle and pelvic girdle are absent, and there is no record of cut marks (Figure 8.35).

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Figure 8.35: Carangidae specimens from Ag-125. NISP= 77

In the case of Clupeidae remains, most of the anatomical regions are absent and were deposited somewhere else. At AG-125 only otic (prootic and pterotic) and occipital

(exoccipital) regions and the vertebral column are present in the sample. No cut marks were observed (Figure 8.36).

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Figure 8.36: Clupeidae specimens from Ag-125. NISP= 87

Figure 8.37: Haemulidae specimens from Ag-125. NISP= 60

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For Haemulidae remains (Figure 8.37), some anatomical regions were absent in the AG-125 sample, specifically those from the branchial arch, pelvic girdle, pectoral and dorsal fins, opercular and investing regions. Perhaps those bones were deposited somewhere else. There is no evidence of cut marks.

Figure 8.38: Sciaenidae specimens from Ag-125. NISP= 34

The pattern observed on Sciaenidae bones (Figure 8.38) suggests that bones from pectoral and dorsal fins, pectoral and pelvic girdle, branchial arch, mandibular, opercular, investing and olfatory regions are absent at AG-125 and perhaps were tossed somewhere else. Just one irregular cut mark was found on an anal spine.

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Figure 8.39: Tetraodontidae specimens from Ag-125. NISP= 63

Finally, the Tetraodontidae´s pattern (Figure 8.39) at AG-125 implies that bones from investing and olfactory regions, pectoral girdle and caudal vertebrae were deposited somewhere else. Just one lateral incision was found on a dentary. Bones with better preservation such as dentary and pre-maxilla are more frequent.

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Chapter 9

Discussion and Conclusions

Introduction

This thesis presents several lines of evidence: zooarchaeological (vertebrate and invertebrate remains), taphonomical (butchering patterns, postdepositional events), anthropic features

(postholes, pits and hearths) and artefactual (pottery, lithic and shell tools) data, which in conjunction; support the hypothesis that Vampiros shelters were places where Pre-

Columbian inhabitants captured and prepare fish. However, the evidence is not sufficiently clear to evaluate the role of Vampiros shelters in the local and regional economy during the

Middle Ceramic Period in the Santa Maria watershed. The contrasting evidence from Sitio

Sierra -a site too close to the coast- suggests that people were also preparing fish products.

There is not sample from an interior village contemporary with Vampiros shelters.

Additional ecological information notes differences through time in fish and shellfish abundance. These differences suggest that during the Vampiros ceramic occupation there were changes from an environment of clear waters and rocky substrates to more estuarine conditions.

In the following sections, I will discuss the zooarchaeological, taphonomical and artefactual evidence.

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Paleoenvironmental Interpretation

Biological and cultural remains were analyzed from samples of both shelters at Cueva de los

Vampiros. Based on 9279 fish remains from both shelters, ca. 80% retained enough morphological features to allow their allocation to some family, genus, and species. Of the

38 families, 82 genera, at least 110 species and 1815 individuals were recognized. Other types of vertebrates were also identified, including, 3 species of mammals, 1 amphibian, 10 species of reptiles and at least 4 species of birds.

As mentioned in the introductory chapters, Central Pacific Panama, specifically

Parita´s Bay, is an area with heterogenic environmental conditions and is affected by climatic and geomorphologic variations. This heterogeneity makes it possible to access several micro- habitats and resources on the estuary. Since the beginning of the Holocene, access to these heterogeneous habitats to obtain marine resources has been limited by changes in coastal geomorphology (Barber 1981; Carvajal Contreras and Hansell 2008; Cooke, et al. 2008).

This transitional, heterogeneous and ephemeral nature of the estuaries is reflected in the Vampiros faunal assemblage. Both molluscs and fish suggest a catchment area along intertidal mudflats close to the river´s mouth. When both Vampiros rockshelters were first occupied by ceramic-using communities around 2300 BP (Macrounit 2b), the surrounding marine ecosystem consisted of clear water and a hard substrate shoreline not far from the coast. This environment at the edge of the estuary supported the presence of green jack

(Caranx caballus), Paloma pompano (Trachinotus paitensis), bonefish (Albula sp.), needle fish

(Tylosurus spp), herrings (Opisthonema), Sierra (Scombermorus sierra), mussels (Mytella), Pacific 357

calico scallops (Argopecten ventricosa) and sea turtles (Eretmochelys imbricate, Lepydochelys cf olivacea,

Chelonia agassizii). According to Cooke (1992a: 25), the fish species mentioned above are occasional visitants of the estuary and move there when there is high visibility, for example, during the end of dry season. Fishing or gathering shells is not a pleasant activity during the rainy season. The alvina and mangrove became muddy areas full of mosquitoes.

Precolumbian inhabitants took up temporary residence along the coast for at least part of the dry season (Cooke and Ranere 1992a, b, c; Linares 1977; Norr 1995; 1996).

Complementary data indicate that during the 500 years of the ceramic occupation, the coastal morphology was constantly changing around Cueva de los Vampiros as a result of coastal progradation in Central Pacific Panama. By the time that the Vampiros shelters were abandoned (upper sediments of macrounit1), there were changes in the species composition abundance. The mollusc and fish remains indicate the middle estuary , near- shore, and ecosystems distributed in mangrove, mudflats, mud sand-rubble , a few sandy beaches, and turbid waters supported molluscs such as grand ark ( Grandiarca grandis), rough littleneck (Protothaca asperrima), moonsnail (Natica unifasciata), pustulose ark (Anadara tuberculosa) and fish such as tete sea-catfish (Ariopsis seemanni), congo sea-catfish (Cathorops),

Kesslers´ sea- catfish (Notarius kessleri), Hachet herring (Ilisha furthii), racer croaker (Ophioscion typicus) and whitefin weakfish (Cynoscion albus). There is poor representation of freshwater fish species and mollusc species that live on sandy beaches or upper estuary.

A stationary trap located 7 kilometers from the mouth of the Santa Maria River reported the following freshwater species in order of abundance: knife fish (Sternopygus dariensis), freshwater catfish (Parauchenipterus amblops), wolf fish(Hoplias microlepsis), panama suckermouth (Hypostomus panamensis), freshwater catfish (Sterisoma panamense) and freshwater

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pimelodid catfish (Rhamdia guatemalensis) (Cooke and Tapia- Rodriguez 1994a) which suggest that Vampiros inhabitants did not exploited freshwater resources or the upper estuary was not part of Vampiros surroundings. There are also low proportions of elasmobranchs.

Finally, there is no evidence of offshore or coral reef species.

Based on the ethology of fish species, Cooke and others (Cooke and Ranere 1999;

1988: 137; Jiménez 1999; Jiménez and Cooke 2001) argued for a technological change during the early ceramic period. People obtained small fish species such as pacifi moonfish (Selene peruviana), brassy grunt (Orthopristis) chalceus thread herring (Opisthonema cf libertate) using fine- meshed gill nets. Cooke also (1992a: 38) suggests that although watercraft is not necessary, it would improve the capture of these fish species using gill nets. This author added that in the case of Pacific moon fish and thread herrings, there is no proof that stationary traps are the appropriate method to capture these clear-water species (Cooke and Ranere 1999: 117).

During the span of at least 300 years of intense fish exploitation at Vampiros

(macrounit 2), I observed a preference for small shoaling species and small species that belong to mangroves such as Pacific moonfish (Selene peruviana), Pacific bumper

(Chloroscombus orqueta), humpback grunt (Ortopristhis chalceus), yellow bobo (Polydactylus opercularis), blue bobo (Polydactylus approximans) and Pacific thread herrings (Opisthonema libertate). Both schooling and mangrove species might have been captured by gill nets.

The other marine species at Vampiros were found on stationary traps located on high salinity waters (20-34 o/00) (Cooke and Tapia- Rodriguez 1994a). This support the idea that

Vampiros people caught the great diversity of fish using traps along intertidal mudflats close to the river´s mouth

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Diversity and equitability of fish and molluscs are low through the entire occupation in both shelters (Macrounit 2b) and increase slightly towards the end of the occupation

(Macrounit 1). Those ecological indexes reflect that people at. Vampiros were focusing on broader environments for shell gathering and fishing several taxa. Fishermen would also take schrimps from their nets and tramps but their remains are not preserved in Vampiros. These strategies are common in the broad spectrum economy of the Middle ceramic period. The number of taxa and species richness indicates, similar to other coastal sites in the Americas, a pattern of extensive exploitation adapted to and dependent on their immediate environments

(Cardera, et al. 2007; Gotz 2008; Reitz and Massucci 2004).

On the other hand, non-fish vertebrate remains indicate that Vampiros inhabitants were not only exploiting and processing fish and gathering molluscs but were also engaged in other local activities in a highly modified coastal landscape. People were birding brown pelican (Pelecanus occidentalis), cormorant (Phalacrocorax cf olivaceus), heron (cf Ardea), gulls

(Laridae) and small passerines. Racoon (Procyon lotor), iguanas (Ctenosaurus similis, Iguana iguana), freshwater turtles (Kinosternon and Trachemys scripta), opossum (Didelphidae), paca

(Agouti paca), and deer (Odocoileus virginianus) were hunted in wooded savannas and terrestrial habitats closer to mangroves. This pattern of exploitation is common to other farming communities identified on the Central Pacific coast (Cooke, et al. 2008)

In general, these aquatic and terrestrial species indicate that Vampiros inhabitants obtained resources near the coast, and they used, perhaps, an uncomplicated (e.g. pebbles for birding) and perishable technology (e.g. intertidal traps, gill nets) to obtain those resources and to create drying plataforms or racks to cure fish. Although marine turtles are

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hunted in Pre-Columbian times, they could be trapped using intertidal traps (Carr 1986;

Cooke and Tapia-Rodríguez 1994a).

Ethnohistorical references indicate that people used cotton nets, line weights, intertidal traps and watercraft to obtain fish:

El manjar mas ordinario de los indios é á que ellos tienen grande afiçion, son los pescados de los rios é de la mar: é son muy diestros en las pesquerías é artificios que se usan.... pescan algunos con caña, de la mesma manera los indios los haçen con varas delgadas é domales é quales convierten para ello, é con cuerdas é volantines é con redes de algodón é muy bien hechas, lo mas continuamente. Y también con corrales é atajos hechos á mano de estacadas en los arreçifes, donde la mar en las costas crece é mengua y en partes á esto apropriadas; y también desde sus canoas ó barcas (Oviedo 1944, III: 210; See also Chapter 3: Footnote 13).

I mentioned before that Central Panamanian archaeological sites lack fishing devices

(net sinkers, hooks) but later in time than Vampiros and contemporary with the late occupation of Sitio Sierra and Cerro Juan Diaz, there is evidence of net weights. Net weights were identified on Pacific sites in the Province of Chiriquí (Linares and Ranere 1980).

Indirect evidence from archaeological, ethnohistorical and ethnoarchaeological records suggest that fish nets and intertidal traps may have been used by the fishers of

Vampiros to capture fish. Perhaps this fishing equipment was made from perishable materials such as wood, cotton, sisal hemp, and bottle gourds or cucurbit gourds (Cooke, et al. 2008; Dickau 2005 ; Griggs 2005; Hart, et al. 2004; Piperno 2000).

Fish poisoning is another fishing method possible used at Vampiros, perhaps to capture anchovies although this fish is scarce on Vampiros samples. Béarez (1998) suggests that Pre-Columbian inhabitants in Ecuador used barbasco (Jacquinia sprucei) or fish poisoning during the first millennium B.C to fish small specimens from the Engrauliade and Clupeidae

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families. This fishing practice is broadly practiced in Latin America (Patiño 1990-1993), and chroniclers such as Oviedo (1535: Libro XIII, Capitulo I) describe the use of fish poisoning in the 16th century:

“Y también usan de cierta hierba que se dice baigua (en lugar de belesa o varbasco), la cual, desmenuzada en el agua, ora sea comiendo della el pescado, o por su propria vertud penetrando el agua, embeódanse los pescados e desde a poco espacio de tiempo se suben sobre el agua, vueltos de espaldas o el vientre para suso, dormidos o atónitos, sin sentido, e los toman a manos en grandísima cantidad. Esta baigua es como bejuco, e picada e majada aprovecha para embarbascar e adormecer el pescado, como he dicho. Pero demás del pescado que así matan en los ríos, toman, de las otras maneras que dije de suso, grande cantidad”.

In Central Panama, Cooke and Jiménez (2008b) recently suggested the use of yam

(Dioscorea spp) as poison to capture freshwater fish at Sitio Sierra.

Similar to fishing technology, there is no evidence of the use of tools to obtain shellfish. Expedient tools for mass collection (i.e. baskets, digging tools) or unselective procurement methods requiring little effort would have been used to collect intertidal shellfish, with the exception of the mobile Pacific calico scallop (Argopecten ventricosus) which either was obtained using an artisanal trawling method attached to small boats or collected during extreme tides called ―aguajes‖. These are monthly spring tides which flooded twice per day for about 5 days (Carvajal Contreras 1998; Martín-Rincón and Rodriguez 2006; B.

Medina, et al. 2007).

The lack of fish equipment in Vampiros and in general in archaeological sites on

Parita Bay contrast with evidence found on archeological sites in Central and South America

(Byrd 1976; Carr 1986; Guzmán and Polaco 2007; Marcus, et al. 1999; Masucci 1995; Reitz and Massucci 2004).

Table 9.1 summarizes the information of four contemporaneous sites which include distance from the sea, date of occupation, predominant aquatic taxa, other vertebrates,

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habitat, probable method of fishing, and artefacts. These sites demonstrate that fishing as part of other subsistence strategies, was a local activity using simple tools (Carvajal

Contreras, et al. 2008; Cooke 1993; Cooke and Jiménez 2004; Cooke, et al. 2007; Cooke and

Ranere 1992a; Cooke and Ranere 1999; Cooke 1994; Cooke 1988, 1992a, 1996; Cooke and

Tapia-Rodríguez 1994a; Hansell 1988; Jiménez 2003; Mayo and Cooke 2005; Mayo 2004).

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Site Distance Date Aquatic taxa Other Habitat Meth. of fishing Artefacts Comments from the Vert. sea(km)

La Mula- vis a vis 3000 - Ariidae, Carangidae, Tetradontidae, Bufo Mangrove, ? Hook and line, No artifacts have Depopulated? At Sarigua coastline 2000 Scianidae, Sphyraenidae. Ostrea spp, marinus, Sandy traps and/ or net been attributed to the end of 1st BP Anadara grandis, Natica unifasciata, Sea turtle, beaches fishing practices millenium B.C. Malea ringens Dayspus?, and tidal due to the Odocoileus flats formation of alvina (salt flats) Cerro Juan 1 km? 2300- Selene peruviana, Opisthonema spp, Ilisha Small Sandy Land-based Line weights in Diaz 1450 furthii, Ariopsis seemani, Guentheridia passerines, beach, fishing Conte Contexts BP formosa and fresh water Sternopygus: duck, intertidal methods/Fine 1650 BP darienensis Iguana, mud, meshed gill nets deer, doves, mangrove egrets. and tidal river Vampiros right on 2300 - Caranx caballus, Chaetodipterus zonatus, Sea Turtle, mud flat, Gill nets/Land- No artifacts have been Cave the 1100 Albula sp, Guentheridia formosa, Odocoyleus mangrove, based attributed to fishing practices coast? BP Ophishonema spp , Iguana, clear water Methods/Wicker Pelican and shallow Traps/poisoning? water / Watercraft. Sitio Sierra 14 km 2300- Fresh water species (P. amblops) and Deer, Near the Gill nets and Net weights and needles from deer 1450 marine species :Sciades dowii, Ariopsis iguana river Land-based metapodials. BP seemanni , Cathorops spp.,Chloroscombrus orqueta,Dormitator latifrons,Selene peruviana and Orthopristis chalceus , Anadara, Ostrea

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Exploitation of Terrestrial and Aquatic Resources

The importance of the different classes of animals at Vampiros based upon the estimated biomass (see chapters six and seven), is a complex matter (Table 9.2). Although mammals and sea turtles provide most of the edible meat in terms of estimated biomass in numbers and most of the NISP and MNI correspond to fish, the transport of cured fish may have caused an underrepresentation of fish bones.

Macrounit 1 Macrounit 2a Macrounit 2b Ctenosaura similis (25200 gr) Chelonia agasizzi (555000gr) Ctenosaura similis (3250 gr) Iguana iguana (9750 gr) Ctenosaura similis (2850 gr) Chelonia agasizzi (245000 Chelonia agasizzi (377000gr) Iguana iguana (9250 gr) gr) Lepydochelys cf olivácea (50000gr) Chiroptera Chiroptera Chiroptera Procyon lotor (1450 gr) Odocoileus virginianus cf Odocoileus Procyon lotor Bufo marinus (1280gr) Phalacrocorax cf punctatus olivieri Sturnella Pelecanus occidentales (700gr) Coragyps atratus Natica unifasciata Natica unifasciata Natica unifasciata Anadara tuberculosa Crassostrea Argopecten ventricosa Prothotaca asperrima Dosinia dunkeri Mytella sp Guentheridia formosa (5380 gr) Guentheridia formosa (15770 gr) Guentheridia formosa Selene peruviana (5140 gr) Selene peruviana (8560 gr) (14760 gr) Sphoerides annulatus (2480 gr) Sphoerides annulatus (7770 gr) Selene peruviana (8200 gr) Notarius kessleri (5880 gr) Polydactylus opercularis (12595 gr) Polydactylus opercularis Orthopristis chalceus (2700 gr) Sciades dowii (17850 gr) 9480 gr) Polydactylus opercularis (4790 gr) Orthopristis chalceus (3140 gr) Sciades dowii (14200 gr) Ilisha furthii (4790 gr) Ilisha furthii (9600 gr) Caranx caballus (8150 gr) Cynoscion albus (13600 gr) Orthopristis chalceus (770 gr) Ilisha furthii (5400 gr) Cynoscion albus (7800 gr)

Table 9.2: Table comparing biomass in some aquatic and terrestrial animals at Vampiros by macrounit

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In general, the estuarine and terrestrial resources recorded for the Macrounit 1 and

Macrounit 2 describe a local and an opportunistic procurement pattern characteristic of this mosaic of habitats. Similar to that described by Cooke (Cooke, et al. 2007; Cooke and Ranere

1992c; 1988), there is a predominance of Pacific moonfish (Selene peruviana) puffer fish

(Sphoeroides annulatus and Guentheridia formosa) and tete-catfish (Ariopsis seemani). The presence of Pacific spadefish (Chaetodipterus zonatus) as well as Panama spadefish (Parapsettus panamensis) and Sierra (Scomberomorus sierra), some anchovies and scallops (Argopecten ventricosa) at the beginning of the ceramic occupation suggest that Vampiros rockshelters might be exploited during the dry season (Cooke 1992a).

A comparison between stratigraphic units indicates that 95% of the sample corresponds to fish, mainly small size fish caught, perhaps, by gill nets or intertidal traps

(Cooke 1988, 1992a). The few sea turtle, mammal, reptile and mollusk individuals may have provided additional meat. The presence of moonsnail opercli (Natica unifasciata) and iguana element distribution and bones with cut marks suggest that the mollusks and black and green iguanas were consumed in Vampiros, rather than being processed and transported elsewhere.

Sea turtles, birds and large-size mammals are represented in Vampiros by low-value elements

(i.e. phalanges, mandibles and carpal elements) suggesting that meaty elements of those animals were discarded or transported for consumption somewhere else.

Shellfish are small prey that may be collected by women and children (Claassen 1998;

Lupo and Schmitt 2002; Moss 1993). Although people collected specimens of mollusks at

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different stages of their growth, small sizes or juvenile specimens were more frequent.

Shellfish were transported and consumed perhaps as a snack or bait at Vampiros. My assumption of the apparent diminution in the sizes of the shells of a marine snail (Natica unifasciata) at Vampiros was not supported by the data (Carvajal Contreras, et al. 2008). In fact, an opposite situation occurs. Smaller individuals of Dunker´s dosinia, pitar venus clam and moonsnail, are found in Macrounit 2b, when the activity of Pre-Columbian fishermen was more intense. The size of invertebrates increases up in Macrounit 1 which coincides with the abandonment of the shelters.

Today sea turtle remains (Cheloniidae), crocodile (Crocodylus acutus), cayman

(Caiman cocodrilus), howler monkeys and whitefaced capuchins (Cebus capucinus) are available at

Parita Bay but according to Cooke and others (Cooke, et al. 2008; Cooke 1992b, 2004a, b;

Linares 1977), are scarce in Parita Bay archaeological middens because of their taboo connotation. Sea turtles‘ iconographic importance is reflected on Pre-Columbian artifacts mainly pottery (Linares 1977:65; Lothrop 1942a). That inference, however, does not fit with the evidence from the Vampiros shelters where there is a fairly high frequency of low-value elements (e.g phalanges, mandibles and carpal elements) suggesting that cured turtle meat was prepared at Vampiros and was transported and/or discarded somewhere else as a part perhaps of a ceremonial or ritual feasting like balseria (Chakraborty 1992; Kirch and O´Day 2003;

Young 1976, 1980a, 1980b).

The information recovered is very important to understanding how native peoples used the middle estuary. In general Vampiros samples reported small (estimated biomass

<400 gr) fish that either grow to a small size, were juvenile specimens or so small (10 to 20 gr)

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that they could have come into Vampiros in the stomachs of carnivorous fish. As I mentioned above, several species of jacks (Carangidae) are associated with clear water, but

Cooke(1992a) mentions that these species enter the estuary under high salinity conditions, to feed (Trachinotus, Oligoplites) or in juvenile stages (Caranx). Other specimens that enter the estuary during high salinity conditions (dry season) are small drums (Sciaenidae) such as the genus Larimus.

Native people in Vampiros captured as well 5 species of snook (Centropomus)

(Robertson and Allen 2006). These are abundant in juvenile stages around the mangrove and coastal lagoons. A few snappers were fished at Vampiros (L. novemfasciatus and L. argentiventris) which are as well abundant as juveniles at estuary but as adults occur over rocky bottoms and are nocturnal predators (Cooke 1992a; Robertson and Allen 2006).

Five species of Corvinas (C. albus, C. squamipinnis, C. praedatorius, C. phoxocephalus,

C.stolzmanni), high fin corvina (Micropogonias altipinnis) and two species of croaker (Bardiella) were identified in the Vampiros´samples. These are estuarine species that lived near river mouths and mangroves (Cooke 1992a).

Several species of marine catfish (Ariidae) were recognized in the samples. The most abundant were the genera Ariopsis and Cathorops (C. tuyra, C. steindachneri, C. furthii, and C. multiradiatus) and few Notarius (N. kessleri, N. cookie, N. lentiginosus), Occidentarius, and Sciades were identified, which live in shallow waters of the estuary. Although some of these catfishes can attain large sizes (i.e Sciades dowi >12 kg), the Ariidae specimens identified at Vampiros are

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relatively small, between 10 to 800 gr, which suggests that Vampiros inhabitants captured juvenile specimens that entered the estuary (Cooke 1992a; Betancur, et al. 2007).

Marine mojarras (family Gerridae; genera Diapterus, Eucinostomus and Eugerres) and white mullet (Mugil curema) were scarce in Vampiros samples, although abundant in the tropical coastal lagoons, which is indicative of changes in the coastal geomorphology at Vampiros

(Robertson and Allen 2006). Several species of grunts (Haemulidae; genera Pomadasys,

Haemulopsis and Orthopristis) are inshore taxa which were found in Vampiros and with a relative small size (150 to 400 gr).

Anchovies (Engraulidae) were present in low numbers from both coarse and fine mesh sieves, which is surprising because of their abundance in Central Pacific Panama. Their low frequency is a result perhaps their having been eaten whole on habitation sites (Cooke and

Jiménez 2004; Cooke, et al. 2004; Cooke and Tapia-Rodríguez 1994a).

Taxa from brackish water, such as the sleepers (Eleotridae), are scarce at Vampiros, contrary to Sitio Sierra, which, as mentioned above, indicates that the pre-Hispanic inhabitants did not exploit the upper estuary.

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Taphonomy and Butchering

My motivation in conducting a taphonomic study was to better understand biases present in the archaeological record as well to know how animal remains accumulate due to human activity or natural processes and their differential preservation within Vampiros deposits.

Natural processes

Several natural processes were distinguished during the analysis of faunal remains. The first marker, abrasion or removal of calcium carbonate in shells as well in vertebrates, was identified in few cases at Vampiros. In a previous article (Carvajal Contreras, et al. 2008), I suggested that abrasion was caused by the effect of salt-aeolian sediments when the shelters were used little or had been abandoned by the Pre-Columbian inhabitants. Other specialists have suggested that this kind of alteration is caused by near-shore wave action or bioeroders like plants (Claassen 1998; Rick, et al. 2006). In the case of Vampiros the abrasion may have been caused by the profuse vegetation that covered the talus before the excavations.

The dissolution of calcium carbonate or the chalky appearance of the shells caused by the presence of water or bioturbation (acid dissolution) was recorded at Vampiros mainly in the uppermost layers (Macrounit 1) and the transition between Macrounit 2a and Macrounit

2b, when at some moment the surface of the shelter was exposed. At the beginning of the occupation of shelters, encrustations or the growth of organisms occur on Argopecten ventricosus’ shells. The worms make the encrustations as well the barnacles when they are alive (Allen and

Costello 1972; Claassen 1998; Rick, et al. 2006)

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Animals perturbed Vampiros deposits. This perturbation was observed during the excavation as burrows made by wasps, crabs, iguanas and rodents, which had displaced vertically faunal and cultural remains, especially in Macrounit 1. These disturbances also affected early contexts and erroneously made it seem that there was marine exploitation during

Preceramic times (i.e., 11,500-7000 BP). Early exploitation of fish resources could have been possible but not with nets and intertidal traps as Vampiros (Carvajal Contreras, et al. 2008;

Cooke 1988; Pearson 2002). In the Vampiros upper layers, unmodified bones from frog, bat, boa, vulture, and owl bones were recovered from both shelters. Most of the perturbed deposits occurred in Macrounit 1 and the uppermost stratigraphic subunits of Macrounit 2.

Probably the unmodified bones represent animals that died of natural causes, that were killed and discarded by the human inhabitants or that nested at these shelters. I did not observe gnawing or tooth marks.

Although the radiocarbon chronology based on charcoal is internally consistent, animal and/or human activities modified the preservation and integrity of the deposits. The completeness of shell specimens, paired bivalves, pottery and articulated remains suggests moments of slow (the transitions between Macrounit 1 and Macrounit 2) and rapid accumulation (Macrounit 2a and 2b). These episodes may represent several discard events or cleaning activities in the living areas and high pre- or post-depositional disturbance when people were trampling, or digging to insert drying racks. As well it may be the result of burrowing activities by frogs, rats and wasps.

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With respect to the occupation, the uppermost layer (Macrounit 1) contained fewer artefacts and had little evidence of ash lenses, hearth and postholes as well animal remains.

The evidence suggests that human activity diminished in this macrounit.

All superimposed anthropogenic features (i.e. postholes, ash floors, charcoal bands and hearths) are contained in Macrounit 2. High density and diversity of faunal remains are coupled with a moderate density of pottery sherds and stone tools.

Both fish and shell remains showed evidence of differential exposure to heat, but it was not possible to determine whether the heat exposure was from cooking/ smoking activities or the result of post-depositional burning. The higher frequency of charred, fragmented and poorly preserved shell and fish material in Vampiros- 2, however, suggests to me that it may have been used either for smoking fish or as a habitational site, whereas

Vampiros- 1 was used more for preparing and drying the fish. . This would be consistent with the fact that Vampiros -2 is shielded from the wind, while Vampiros- 1 is exposed to the wind (see Figure 9.1) (Carvajal Contreras, et al. 2008).

Dr. George Pearson, during his excavation of Vampiros-2 trying to recover

Paleoindian remains, recorded several features. One of these was (Figure 9.1) a deep hearth, approximately 1.2 X 1m, surrounded by postholes (indicated by wood sticks) that could have been associated with smoking fish. Processing sites documented archaeologically and ethnographically show some similarities with Vampiros. These similarities are: fish bones , processing facilities (e.g. postholes, hearths), location near marine resources, water supply (e.g.

Santa Maria river), salt resources (alvina), low proportions of other animal remains, cut marks,

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a shift of element representation (more cranial than postcranial bones) (Barrett 1997; Gifford-

Gonzalez, et al. 1999; Høtje 2006; Stewart and Gifford-Gonzalez 1994).

Figure 9.1: Vampiros-2: hearth surrounded by postholes (toothpicks) by Pearson.

Butchering

Comparisons of the butchering patterns of the coetaneous Vampiros and Sitio Sierra and the later site AG-125 proved informative. I had predicted that Vampiros and AG-125 would share a similar pattern since both sites were located on the coast at the time of their respective occupations. I also predicted that the pattern at Sitio Sierra would be different from those two sites because Sitio Sierra was located far from the coast, assuming that traveling down the river by boat to obtain marine fish was a dangerous enterprise and that it is more plausible that the marine resources were obtained by internal trade (Cooke and Jiménez 2004; Cooke, et al.

2008; Cooke and Ranere 1992a; Cooke and Ranere 1999; Cooke and Sanchez 2001).

It was difficult to establish a standardized proportion of the fish bone elements (e.g. anatomical regions, cranial vs. post cranial) since they vary by species. However, I compared

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the element distribution and proportions of Ariidae, Carangidae, Clupeidae, Haemulidae,

Sciaenidae and Tetraodontidae for the three sites, based on the cured-fish model.

This model is challenged somehow by Cueva de los Vampiros, Sitio Sierra and AG.-

125 data. The pattern for the Ariidae, Carangidae, Clupeidae, Haemulidae, Sciaenidae and

Tetraodontidae families indicates there are overabundances of cranial bones in comparison with appendicular elements at Vampiros, Sitio Sierra and AG-125, which suggests that at all three sites fish were being processed.

An exception is for the Ariidae at AG-125 that shows an overabundance of axial elements; there are more pectoral fin and dorsal fin bones represented in Ag-125 than Vampiros and

Sitio Sierra. It may be interpreted that the skull and trunk from Ag-125 were transported somewhere else.

It seems that Sitio Sierra, Ag-125 and Vampiros have more similarities than I expected. The missing or less frequent bones on those sites for all families are the pelvic girdle, opercular region, and branchial arch which suggests that they were deposited somewhere else or possibly destroyed in situ. In Sitio Sierra the Carangidae, Clupeidae, Haemulidae and Sciaenidae families have more upper vertebrae than Vampiros and Ag-125 which are indicative of stockfish processing.

The three sites share a similar Sciaenidae‘ general distribution: the missing trunk portion, which was transported somewhere else. Although in the Vampiros sample there were discarded more bones from the anal and pectoral fins, Sitio Sierra and Vampiros also share a

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similar Carangidae´s pattern: a lower frequency of bones from the mid-section, opercular and pelvic girdle which were discard or transported to another location.

In the case of the Clupeidae family, most of the skull regions are underrepresented with the exception of the occipital and otic. The vertebral series indicate that AG-125, Sitio

Sierra and Vampiros lacked of upper vertebrae (thoracic) and precaudal vertebrae. That profile suggests that the whole animal was transported somewhere else.

The cured model also is challenged by the presence of the cleithra from the appendicular region. In this model low frequency of cleithra is indicative of a processing site, whereas their high frequency suggests a distribution of a habitation site (Amundsen 2008;

Krivogorskaya, et al. 2005; Krivogorskaya., et al. 2005; Perdikaris 1999). Vampiros, Sitio Sierra and Ag-125, show high frequencies of cleithra, mostly in fragmented state. The cleithra presence is congruent with the model proposed by Zohar and colleagues (Zohar and Cooke

1997; Zohar, et al. 2001b) where for both small and large sized fish the cleithra is the most damaged bone from butchering.

With respect to Ag-125, its proximity to the sea suggests that it should share a similar element distribution to Vampiros. At this site, there are more cranial than postcranial bones and most of the sample comprises bones from fins and the pectoral girdle, suggesting that the people were gutting fish. However, some missing bones either were deposited somewhere else or were differentially affected by other agents. Since AG-125 was an open site located on the coast, other taphonomical factors were involved in the preservation of fish remains (Stahl

1995, 2000).

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The three sites show abundant presence of bones belonging to the vertebral column, pectoral girdle and lateral region, which are the most robust elements. Elements from the neurocranium such as the olfactory, investing and opercular regions are the least frequent, due probably to their extreme fragility and their failure to survive butchering, dropping and trampling or their differential transport. Postcranial bones such as vertebrae are more robust and survived better.

As mentioned in previous sections, Vampiros assemblages display more cut marks than at the other two sites, which reinforces the hypothesis of its function as a processing site.

It is difficult to evaluate whether smaller animals were treated differently than bigger specimens as Zohar and colleagues have suggested (Zohar and Cooke 1997). Most of the sample from Vampiros rockshelters corresponds to small size animals. The fish size information was not available for Sitio Sierra and Ag-125. Despite this inconvenience, small specimens were processed at Vampiros and Sitio Sierra. Specifically whole skulls and fully articulated specimens were found. These articulated fragments partially corroborate Zohar´s hypothesis of differential butchering methods according to the size of the specimen (method

1) in which some fish are butchered and perhaps cured keeping the head intact.

In general, cut marks were located on the midline side, which indicates the action of removing gills and entrails for salting and drying in Vampiros, and more lateral marks were found in Sitio Sierra, which could suggest the removing of the flesh. On the other hand,

Vampiros, Sitio Sierra and Ag-125 fracture patterns show that the best preserved elements are the vertebrae. Damaged skull bones were found at Vampiros, Sitio Sierra and Ag-125,

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especially the cleithra fragments of which were discarded, suggesting that perhaps the specimens were dorsally split along skull and vertebral column for splitting the fish for salting and/or drying (medial and lateral cut marks and several fracture portions) and removing guts.

The presence of first dorsal spines and anal spine and anal pterygiophores suggests these bones, were removed to reduce injuries during the handling of the specimens. Cut mark and fracture locations imply cutting or chopping to decapitate the animal (supracleithrum, coracoids, cleithrum, vertebral complex), to remove gills (epihyal, ceratohyal, opercle), fins

(pterygiophores, spines) and tail (caudal vertebrae). The cut mark pattern observed implies that most of the cuts were transversal and made with fine instruments (La Mula points may have produced fine lines) or massive tools (polished axes may have produced chopped, crushed or impact marks).

These cut marks and fractures would be entirely consistent with a site where people processed fish for curing nor for filleting. Vampiros cut marks contrast with the paucity of cut marks, found at Sitio Sierra and AG-125, which implies that fish perhaps were not processsed only for curing but for consumption at those sites, but this remain inconclusive as the pattern of fractures show similarities between Vampiros and Sitio Sierra. Surprisinly cut marks were also identified in the other vertabrates analyzed at Vampiros which suggests that people were prepairing other animal products: sawing marks on raccon, Ardea and Strigidae, spiral fractures on deer and sea turtle (Chelonia agasizzi), and incisions on Iguana bones.

People at Vampiros were perhaps scaling fish. Carangidae´s scales were recovered in the samples, but I cannot dismiss the possibility that the same activity was carried out at Sitio

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Sierra and Ag-125 where no scales were identified. This situation could be a product of differential preservation (Stewart and Gifford-Gonzalez 1994; Zohar 2003; Zohar and Cooke

1997; Zohar, et al 2001).

The relative representation of the vertebral column at Vampiros indicates that most of the discarded vertebrae belong to the section adjacent to the head or thoracic vertebrae

(Ariidae, Polynemidae, and Tetraodontidae) and the section adjacent to the tail or caudal vertebrae (Sciaenidae and Clupeidae). This pattern is shared as well by Sitio Sierra and partially by AG-125. This could be mean that the spine was divided and the upper section was thrown as individual units with the head during the butchering process. The upper caudal vertebrae and most of the precaudal vertebrae are absent at Vampiros, AG-125 and Sitio

Sierra which were left perhaps in the exported products.

To sum up, there are two patterns. The first pattern suggests that smaller individuals were being transported almost complete (Clupeidae). The second pattern is a mix, suggesting that small individuals were partially discarded and that the trunk portions were transported, while large individuals were partially butchered and the trunk was transported elsewhere as well ( Ariidae, Carangidae, Haemulidae, Sciaenidae and Tetraodontidae). The Vampiros assemblage has a similar frequency of bones from the pectoral girdle, the branchial region, and the hyoid arch to the other two sites. Those elements, that Tourunenn (2008) called ―throat‖, were removed and discarded in Vampiros and Sitio Sierra. This information suggests that people at Vampiros were scaling, gutting and cutting fish.

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Vampiros stands out because of the abundance of Tetraodontidae (puffer fish) bones in its deposits. Vampiros and AG-125 have similar distributions of the lateral and mandibular regions, but Ag-125 lacks sections of the neurocranium and lower vertebrae.

Based on element profiles, cut marks and fractures, Pre-Columbian inhabitants at Vampiros were perhaps removing the skin, guts, tail and head of the Tetraodontidae animals and popping-out two dorsal fillets as fractures and cut marks on supracleithrum, dentary, premaxila and vertebrae are found. The zooarchaeological literature (Cooke, et al. 2008;

Cooke and Ranere 1992a; Cooke and Ranere 1999; Cooke 1988, 2001) and the present evidence here note the high frequency of puffer fish at Vampiros in contrast with other sites in Central Pacific Panama such as Cerro Mangote, Aguadulce Shelter, Monagrillo, Sitio Sierra,

AG-125, and Cerro Juan Diaz, with samples dated between 5000 BC to 1300 AD, with the exception of the contemporaneous site La Mula-Sarigua and later site La

Pitahaya(GranChiriqí) (Cooke and Jiménez 2004; Wing 1980).

The comparison between Vampiros and Sitio Sierra´s Tetraodontidae assemblages may be explained as exchange of marine resources such as puffer fish, but for the other families the evidence is inconclusive. Several authors point out from Cerro Mangote and Sitio Sierra´s archaeological and isotopic data that different groups on the Central Pacific coast of Panama were either moving seasonally between coastal and inland sites or as a single group were occupying both coastal and inland sites, and exchanging resources between them (Cooke, et al.

2008; Cooke 2005; Norr 1995, 1996).

Several factors would be also involved in differential preservation of each anatomical element. I focused on mainly butchering as a result of the processing of fish but scavenging,

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trampling, sediment chemistry and other processes that occur in human occupation sites would also affect the preservation of these remains. That could be a possible explanation for some underrepresented areas such as the opercular, branchial arch and investing bones.

Further studies on bone density of these tropical families would clarify this matter.

Finally , from a methodological point of view, although the skeletal element distribution, MNI and MAU were good tools to observe butchering patterns, these are somehow disturbed by the better preservation of robust elements in all families (i.e. dentary, premaxilla, anal pterygipohore, pectoral spines, vertebrae ) that give the impression of being the most abundant elements overall. Robust and fragile elements are distinctive when recovered intact in Ariidae, Carangidae, Haemulidae, Polynemidae, Pristigasteridae and

Tetraodontidae, but like the other bones of the following families, they are quite fragile and easily destroyed by normally occurring taphonomic forces: skull parts of Belonidae,

Engraulidae, Eleotridae, Clupeidae and Mugilidae. There were also difficulties in assigning

MAU and MNI values, especially vertebrae, since each species has a different number.

Exchange of Fish and Specialization in Panama as seen from Vampiros Rockshelters.

The Vampiros evidence is not enough to understand the exchange of seafood resources or how some communities specialized in some economic activities, in this case coastal exploitation in comparison to Sitio Sierra and AG-125, or to recognize Vampiros as a specialized site for producing preserved fish per se. In the Americas zooarchaeological evidence for chiefdoms and state-level societies has been used to examine the exchange of food resources, either terrestrial (i.e. guinea pig, camelids, deer and Muscovy duck) or marine 380

resources (i.e. thorn oyster), as part of the political economy of these societies (deFrance 2009;

Emery 2004; Marcos 1986; Paulseen 1974; Reitz and Massucci 2004; Sandweiss 1996; Stahl

1995). Several scholars have argued that coastal resources have been the basis for specialization and the development of complex societies (deFrance 2001; Moseley 1992a;

Sandweiss, et al. 1999; Sandweiss, et al. 1999; Sandweiss, et al. 1998).

Studies of the exchange of fish and of communities specialized in curing and processing fish are scarce (Carr 1987; Cooke 1988; Guzmán 2008). These are limited to ethnoarchaeological evidence (Stewart and Gifford-Gonzalez 1994; Zohar 2003; Zohar and

Cooke 1997; Zohar, et al 2001), and the dichotomy between coastal versus inland sites and their element distribution and cut mark distribution (Amundsen 2008; Carr 1986, 1987; Cooke

1988; deFrance 2009; Gotz 2008; Krivogorskaya, et al. 2005; Krivogorskaya., et al. 2005;

Masson and Peraza 2008; Perdikaris 1999; Reitz and Massucci 2004). Studies of the exchange of fish are limited as well as to indirect evidence from the archaeological context such as fishing equipment (Marcus 1987; Marcus, et al. 1999), historical records (Barrett 1997;

Brewer and Friedman 1989; Høtje 2006) and diversity of species at the processing site (Bearez

1996; Reitz and Massucci 2004; Voorhies 2004). The exchange and curing of fish remains is also tied with the use and production of salt (Griggs 2005; Mock 1994).

It is assumed that low animal species diversity and abundance is associated with a special purpose site (Guzmán 2008; Stewart and Gifford-Gonzalez 1994; Zohar 2003; Zohar, et al. 2001b). As observe in the following Table, Vampiros has an outstanding diversity in comparison with the other two sites. This great diversity of fish taxa accounted in Vampiros is result of the inhabitants collecting all the fish caught in their nets and /or traps.

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Site Families Genera Species MNI S. Size Vampiros 38 82 110 1815 9279

Sitio 13 34 63 408 15549 Sierra

AG-125 22 42 54 112 622

Table 9.3: Summary of Vampiros, Sitio Sierra and Ag-125

The problem of food trade is modified in the case of Vampires by several aspects: social organization, seasonal mobility and the pattern of settlement. Following Sahlins1972), Polanyi

(1957) and Renfrew (1975), I propose the following scenarios. One possible scenario is that people at Sitio Sierra did not consume puffer fish, which would account for the absence of bones from this family in its deposits. Vampiros and Sitio Sierra were autonomous and self- sufficient groups and fished, butchered and cured their own marine resources (Renfrew´s internal trade direct access model). The Vampiros puffer fish assemblage shows cut marks that are related with decapitation and removing skin (fine incisions on the supracleithra) and guts.

The other possible scenario, if Vampiros was part of the same social unit as

Sitio Sierra, is that Vampiros was a fishing station -similar to deer hunting reserves or cotos— that processed puffer fish and/or turtle and exported the meat as a delicacy during a ceremony like balseria to Sitio Sierra during the dry season (Renfrew´s home base reciprocity or down the line models).

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Puffer fish have been consumed as fugu, sashimi, or in salad in Asian and African countries for centuries. The attraction of this exotic food, potentially poisonous, is the experience of a numbness on the lips and tongue while eating and shortly thereafter. The puffer fish neurotoxin may also cause a zombie-like state of apparent death (Davis 1997;

Tanaka 2008; Wolinsky, et al. 2005).

It should not be surprising that puffer fish were consumed in Central Panama not only in Vampiros but at La Mula-Sarigua and a site on the coast of Chiriqui -La Pitahaya. The use of toxic foods by early hunter-gathers and early farmers before and after the emergence of food production, especially plants such as certain nuts and tubers, has been recorded in the

Americas and worldwide. Several authors argue the time-consuming processing of toxic resources is worth the effort when compared with the high productivity of those plants and animals(Grivetti 1981; Wilson 1997). Animal products such as frogs, stingrays, coral, shellfish and puffer fish have been used in the past and today for dietary and religious purposes in

Panama, Haiti, Egypt and Guatemala (Brewer 1989; Brewer and Friedman 1989; Cooke 1989;

Davis 1997; Glowacki 2005; Maxwell 2000). In particular, puffer fish (Tetraodontidae family) are highly appreciated today, not only in Asian countries as fugu, but in Central and South

America and Mexico as well where there are farms dedicated to its production. This family contains tetraodoxin, a venom without antidote, which is concentrated in muscle, gonads, skin and viscera. It causes death without proper preparation (Bearez 1996; Chavez Sanchez, et al.

2008; García-Ortega, et al. 2002; Guzmán 2008; Nunez-Vazquez, et al. 2000; Stahl 2003).

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The remains form other families suggest that Vampiros, Sitio Sierra and Ag-125 were processing fish. The pattern of fragmentation of the cleithrum on the three sites suggests that native people were cutting the head for salting and drying fish.

The other possible scenario assumes that Pre-Columbian fishers from Vampiros, Ag-

125 and Sitio Sierra were eating their own catch as fresh fish (Renfrew´s internal trade direct access model). Therefore, middens would have a balance of skeletal elements more similar to that found in live fish. I do not think that these three sites fit this situation.

In this temporal context and based on the evidence, I consider that Vampiros is an early site for processing fish during the dry season. I think Vampiros and Sitio Sierra, both probably belonged to the same political unit. They were self-sufficient and autonomous in terms of food-stuff and probably caught and prepared the fish themselves for their own consumption and could exchange some preserved fish with their relatives in the Central foothills. I cannot prove or dismiss that Vampiros was the supplier of Sitio Sierra´s marine fish.

This assumption, with respect to Vampiros, is partly supported by the data. The following lines of evidence clarify it: 1) the associated features related to curing and/or smoking fish, 2) the relative abundance of cranial versus appendicular bones, 3) the relative representation of different sections of the vertebral column compared with cranial elements;

4) the types of bones in articulated groups; 5) presence of scales, 6) the distribution of cut marks. However the pattern observed at Sitio Sierra, also observed by Cooke(1988), is less conclusive. The distribution of elements indicates that the Pre-Columbian inhabitants obtained some marine fish (Tetraodontidae) from a site similar to Vampiros, but other

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resources may have been obtained by them (the remaining families). It is possible that differences and/or similarities between fish families in Vampiros and Sitio Sierra´s assemblages are caused by the discarding pattern in Sierra, a domestic context (i.e. midden), and by its distance to the sea at the time of occupation. Either that or groups moved from the coast to the interior in a trashumance pattern of seasonal rounds (Cooke 2005; Cooke and

Ranere 1992a, 1999; Ranere and Hansell 1978).

The antecedents of trade of animal products can be seen in Panama during Preceramic times (ca. 5000 B.C). A site located on the Pacific coastline, Cerro Mangote, has evidence of manatee bones from the Caribbean. The existence of inland contemporaneous sites such as

Cueva de los Ladrones, that also contain fish remains, suggests that these resources were obtained from a coastal site by exchange with local communities (Cooke, et al. 2008; Cooke

2005).

The Vampiros site and contemporaneous sites in Central Pacific Panama (la Mula-

Sarigua, Cerro Juan Díaz, Bucaro and Sitio Sierra) during ca. 200a.C to A.D. 200 were immersed in a period which saw the emergence of village agricultural life and social differentiation based on sex and occupation (egalitarian society) as the mortuary goods recovered from burials are almost all utilitarian (Briggs 1989; Cooke 1984; Cooke and Ranere

1992b; Hansell 1988; Ichon 1980; Isaza 1993). These sedentary communities were nucleated on ecotonal locations which gave them differential access to local lithic, plant and animal resources and increased residential stability. These communities might have started to develop territories around these rich and diverse areas (Carvajal Contreras and Hansell 2008;

Cooke, et al. 2008; Hansell 1988; Jackson 1983). That they developed a broad spectrum

385

economy during the Middle ceramic period implies self-sufficiency in terms of subsistence activities by the household unit, but fishing and shell-fishing may have been carried out through collective investment of several domestic units during the dry season (Cooke, et al.

2008; Cooke and Ranere 1992b; Cooke and Sanchez Herrera 1997; Haller 2004; Hansell 1988;

Isaza-Aizpurua 2007; Norr 1995). By this time, specialists started to emerge, as is evident in the chipped stone tool production which employed standardized techniques of manufacture, and a regional exchange network, perhaps organized along kinship lines, between coastal populations and inland communities for goods such as polished axes, chipped-stone tools, dried and salted fish, and possibly salt (Cooke and Ranere 1992b; Haller 2004; Hansell 1988;

Isaza-Aizpurua 1993; Isaza-Aizpurua 2007).

Comparisons

There are obstacles to comparing butchering practices of prehistoric people in Panama and Lower Central America. First, there is little zooarchaeological research addressing the distribution of elements; instead, research focuses on changes of fish frequencies over time, indicating access to several habitats and fishing techniques (Bearez 1996; Béarez 1998;

Gutierrez 1997; Hamblin 1984; Julian Kerbis 1980; Reitz and Massucci 2004; Rewniak 2006;

Stahl 2003; Wing 1980). Second such investigations face a diversity of fish species in the

Neotropics and the consequent difference in number of bones per species (Robertson and

Allen 2006). Third, there are few ethnoarchaeological works addressing fish butchering patterns (Zohar and Cooke 1997). Finally, there is no standard to record the anatomical parts and discern patterns of transportation and disposal of fish remains in each zooarchaeological work (Carr 1987; Cooke 1988; Creamer 1983; Guzmán 2008; Masson and Peraza 2008).

386

A preliminary attempt to investigate the proportional distribution of body parts in

Panama was carried out by Cooke (1988) comparing samples from Cerro Mangote,

Aguadulce, Sitio Sierra, Cueva de los Vampiros and La Mula-Sarigua dated 5000 B.C. to A.D.

500. This study is based on percentage of total fish sample (NISP %) of vertebral column, head, spine, rays, pterygiophores and otoliths on samples not completely analyzed and number of charred bones.

The main conclusion of the fish element distribution between Vampiros and Sitio

Sierra is that Vampiros was a fishing station and Sitio Sierra obtained marine fish through trade and obtained locally fresh water taxa. There is a higher proportion of Ariidae´s skulls, vertebrae and spines in Sitio Sierra than Vampiros. There are frequent Tetraodontidae´s skull bones at Vampiros and La Mula- Sarigua than the other sites. Carangidae´s skull and vertebrae are more frequent at Vampiros than Sitio Sierra. Finally, Clupeidae vertebrae are more frequent in Sitio Sierra than Vampiros.

This exchange pattern between coastal and inland communities is repeated in later times in Central Panama. Ilean Isaza Aizpurua (2007) during a regional survey along the La

Villa River discovered a fishing village (LS-31). This site was situated along an active coastline and occupied between the Tonosí phase (Middle Ceramic B) the −Conte phase (Late Ceramic

A), ca A.D.550 and 700. She suggests that this coastal site was used for processing and redistributing fish (food) and shell (raw material for ornaments). Isaza further suggests that

Monagrillo, La Mula-Satigua and LS-31 share some characteristics, mainly their location and species composition, which makes these villages candidates for processing and redistribution sites. However, Isaza did not carry out the kind of quantitative analysis (i.e. element distribution) that has been presented in her thesis. 387

Winifred Creamer (1983) identified fish body parts (NISP) to detect butchering patterns that would explain food exchange on two islands in the Gulf of Nicoya on the Pacific coast of Costa Rica: Chira and San Lucas. Her working hypothesis was that offsite butchering is characterized by bones associated with meat cuts without heads. She pointed out that fish elements are not good indicators of exchange because most of the unidentified portion could account for ―missing‖ bones of either vertebrae or skull bones. Second, modern fishermen in

Costa Rica are ―neither filleting nor removing heads‖, which is similar to the method described by Zohar. Finally, she suggests that dried fish trade would be represented only on inland sites.

Two Mexican sites on the Pacific Coast, Huatabampo, occupied between 177 BC and

AD 700–900, and San Felipe Aztatán dated, AD 900 to 1521, provide an interesting comparison. At Huatabampo inhabitants preferred mullets (Mugil sp), croakers (Pomadasys sp) and sea catfish (Ariopsis seemani, Cathorops sp) from the mouth of a coastal lagoon, whereas at

San Felipe Aztatán´s inhabitants favoured sea catfish (Ariopsis seemani) and cichlids (Cichlasoma beani) from the inner side of a lagoon. Based on high NISP values from Huatabampo and San

Felipe Aztatán the assemblages, Guzman mentions that fish ―show the presence of head, vertebral, and fin bones, with a better representation of the anterior half of the body. Some burned bones are evidence of immediate consumption (if roasted) or preservation for later consumption (if smoked). This pattern suggests that the inhabitants processed and consumed the whole animal in situ ―(Guzmán 2008: 42). Significantly there is an absence of mandibular and hyomandibular arches and gill apparatus.

A preliminary report from the Maya site of Isla Cerritos by Helen Carr (1987) indicated that Pre-Columbian inhabitants exploited mainly catfishes (Siluriformes), porgies 388

(Sparidae) and drums (Sciaemidae). Carr compared the proportion (NISP %) of cranial

(syncranium) and postcranial (vertebrae and fin and elements) at Isla Cerritos with Cozumel

Island, Chichén Itzá and Cerros. She argued that Isla Cerritos was a site where fish was processed to export with 53% of cranial remains, whereas Cozumel Island, Chichén Itzá and

Cerros, the other sites, presented high percentages of postcranial bones (63%, 53%, and 51% respectively). However, Carr is cautious in her interpretation since fine- screen materials could reverse the situation at Isla Cerritos (Carr 1986: 21).

Zooarchaeological evidence at the Postclassic Maya city of Mayapán indicates that certain fish marine species such as catfish (Ariopsis felis), tarpon (Megalops atlanticus), snappers (Lutjanidae) and bass (Serranidae) were obtained through trade (Masson and Peraza 2008). These authors, based on the cranial, fins and vertebrae frequencies (NISP %) of catfish and non- catfish specimens, suggest that the higher ratio of vertebra to cranial parts of non-catfish may indicate the procurement of postcranial salted non- catfish carcasses (with vertebrae and spines intact) from coastal localities, whereas the higher frequencies of catfish skulls indicate that Pre-

Columbian residents obtained catfish directly or they were procured through trade as preserved whole fish.

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Conclusions and Future Research

To conclude, this research accomplished the following points:

1. Vampiros shelters were used intensively for exploiting resources of the middle estuary

and the mangrove from ca 2300-1800 rcya (The Middle Ceramic); consistent with prior

models for coastal progradation.

2. One hundred ten species of fish have so far been identified. All were marine inshore

caught by intertidal trap or gill nets. Fish and mollusk species that prefer clear water

columns (green jack, Pacific calico scallop etc) were more frequent in lower levels. There

was no substantial use of freshwater species. Estuarine species adapted to more turbid

water conditions were frequent in upper levels.

3. There is physical evidence for fish butchering (scaling, gutting, cutting) at Vampiros.

The evidence is congruent to butcher fish for salting and drying (Zohar and Cooke

1997).

4. Differential distribution of burnt and fragmented bones, shells and artefacts suggests

windward and leeward shelters were used for different activities. Vampiros-1 was used to

preserve fish and Vampiros-2 was used as a habitation or smoking fish.

5. Sea turtle bones (mandibles and phalanges) were considerably more abundant in

Vampiros than at other middens at Parita Bay sites. Perhaps Vampiros shelters produced

smoked turtle meat to export to inland sites for feasting.

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6. Remains of terrestrial vertebrates were more diverse and abundant than predicted,

suggesting other economic activities were carried out at Vampiros.

7. Puffer fish were consumed and prepared. Vampiros was a site that specialized to

remove puffer fish´s skin and visceras and to export the meat as a delicacy to sites

further inland.

8. Small fish were transported almost complete (Clupeidae), while the other pattern is a

mixture suggesting that small specimens were partially discarded and trunk portions

were transported and large individuals were partially butchered and the meat was

transported elsewhere (Ariidae, Carangidae, Haemulidae, Sciaenidae and Tetraodontidae).

9. Compared to other sites (Sitio Sierra and AG-125), Vampiros‘ role in the local and

regional economy is not clear. The three sites had bones from the pectoral girdle, the

branchial region, and the hyoid arch. This distribution suggests that those sites were self-

sufficient and were gutting and preparing their own fish resources. However, lateral cut

marks on Sitio Sierra´s fish sample suggest other type of preparation for consumption.

The past thirty years in the Central Pacific Panama has under gone a progressive and

steady increase in archaeological and zoo-archaeological research, specifically within

icthioarchaeology (Carvajal Contreras, et al. 2008; Cooke 1993; Cooke and Jiménez 2004;

Cooke and Jiménez 2008a, b; Cooke, et al. 2007, 2008; Cooke 1988, 1992a; Cooke and

Tapia-Rodríguez 1994a, b; Jiménez 1999; Jiménez and Cooke 2001; Zohar and Cooke

1997). Throughout the region numerous researchers have contributed an enormous

391

amount of work to this niche alone, particularly the study of identification, ecology,

subsistence strategies, technology.

This dissertation offered information regarding taphonomy, fish butchering practices and early development of an exchange system between coastal and inland site that include fish and turtle. To understand fully if Pre-Columbian inhabitants butchered fish according to size, it is not clear at Vampiros shelters since most of fish caught by pre-Columbian inhabitants correspond to small size specimens. If a full analysis of the fish remains from the column samples of Vampiros were carried out, we would have a better understanding of butchering practices by species and size. I propose for future research to demonstrate that the exchange of cured fish changed as social organization changed from egalitarian to hierarchical.

Furthermore, excavating inland site occupied during the Middle and Late Ceramic with good faunal remains preservation would provide data to observe how a consumer site looks like not only in terms of the faunal but pottery and lithic assemblages as well.

392

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429

APPENDIX 1 Ariidae skeletal elements in a complete fish.

Anatomic region/ Expected no. Skeletal element Olfactory Region (n=12) Ethmoid 1 Prefrontal 2 Supraethmoid 1 Vomer 1 Parasphenoid 1 Alisphenoid 2 Total orbital Series 4 Occipital Region (n=5) Exoccipital 2 Supraoccipital 1 Basioccipital 1 Otic Region (n=16) Prootic 2 Pterotic 2 Sphenotic 2 Opistohotic 2 Epiotic 2 Otoliths no. 6 Investing Region (n=8) Nasal 2 Frontal 2 Parietal. 2 Scale bone 2 Lateral region (n=14) Premaxilla 2 Maxilla 2 Dentary 2 Articular&Angular 4 Preopercle 2 Lacrymal 2 Mandibular arch (n=8) Mesopterygoid 2 Metapterygoid 2 Quadrate 2 Palatine 2 Hyoid region (n=11) Hyomandibular 2 Interhyal 2 Epihyal 2 Ceratohyal 2 Hypohyal 2 Glossihyal 1 Opercular Series (n=8) Branchiostegal ray 4 Opercular 2 Interopercular 2

430

APPENDIX 1- cont'd

Anatomic region/ Expected no. Skeletal element- Branchial Arch (n=22) Epibranchial 6 Ceratobranchial 8 Hypobranchial 4 Basibranchial 2 Pharyngobranchial 2 Urohyal 1 Pelvic Girdle (n=2) Pelvis (basipterygium) 2 Pectoral Girdle (n=10) Cleithrum 2 Scapula 2 Coracoid 2 Posttemporal 2 Supracleithrum 2 Fins (n=4) 1st dorsal spine 1 Pterygiophore dorsal 1 Pectoral spines 2 Vertebra total no. (46) Atlas 1 Fused vertebrae (included tripus) 1 Precaudal 10 Caudal 32 Penultimate 1 Ultimate 1

431

APPENDIX 2-Haemulidae, skeletal elements in a complete fish.

Anatomic region/ Expected no. Skeletal element Olfactory Region (n=17) Ethmoid 1 Vomer 1 Prefrontal 2 Alisphenoid 2 Parasphenoid 1 Orbital series 10

Occipital region (n=4) Supraoccipital 1 Exoccipital 2 Basioccipital 1 Otic Region (n=16) Epiotic 2 Opisthotic 2 Prootic 2 Pterotic 2 Sphenotic 2 Otolith 6 Investing region (n=6) Nasal 2 Frontal 2 Parietal 2 Lateral region (n=12) Premaxilla 2 Maxilla 2 Dentary 2 Articular&Angular 4 Preopercle 2 Mandibular region (n=8) Ectopterygoid 2 Metapterygoid 2 Quadrate 2 Palatine 2 Hyoid region (n=12) Hyomandibular 2 Symplectic 2 Interhyal 2 Epihyal 2 Ceratohyal 2 Hypohyal 2 basihyal 1 Anatomic region/ Expected no. Skeletal element- Opercular Series (n=20) Opercular 2 Interopercular 2 Subopercular 2 Branchiostegal ray 14 Branchial Arch (n=41) 432

pharyngial plate 4 Epibranchial 6 Ceratobranchial 8 Hypobranchial 8 Basibranchial 8 Pharyngobranch 6 Urohyal 1

Pelvic Girdle (n=4) Pelvis 2 Pelvic Spines 2 Pectoral Girdle (n=12) Cleithrum 2 Coracoid 2 Postcleithrum 2 Scapula 2 Supracleithrum 2 Posttemporal 2 Vertebrae n=26 Atlas 1 Thoracic 3 Precaudal 6 Caudal 14 Penultimate 1 Ultimate 1 Fin (n=4)

1st anal pterygiophore. 1 Anal Spine 3 Dorsal spine 1 Pectoral spine 1

433

APPENDIX 3- Carangidae skeletal elements in a complete fish.

Anatomic region/ Expected no. Skeletal element-Cranial Olfactory Region (n=17) Lateral Ethmoid 2 Vomer 1 Prefrontal 2 Supraethmoid 1 Alisphenoid 2 Parasphenoid 1 Total orbital Series 8 Occipital Region (n=4) Basioccipital 1 Exoccipital 2 Supraoccipital 1 Otic Region (n=16) Opisthotic 2 Epiotic 2 Prootic 2 Pterotic 2 Sphenotic 2 Otolith 6 Investing Bones Region (n=6) Parietal 2 Nasal 2 Frontal 2 Lateral region (n=14) Premaxilla 2 Maxilla 2 Dentary 2 Articular&Angular 4 Preopercle 2 Lachrymal 2 Mandibular region (n=8) Palatine 2 Ectopterygoid 2 Metapterygoid 2 Quadrate 2 Hyoid region (n=12) Hyomandibular 2 Symplectic 2 Interhyal 2 Epihyal 2 Ceratohyal 2 Hypohyal 2

434

APPENDIX-3 cont'd

Anatomic region/ Expected no. Skeletal element- Opercular Series (n=16) Branchiostegal ray 10 Opercular 2 Interopercular 2 Subopercular 2 Branchial Arch (n=41) pharyngeal plate 4 Epibranchial 6 Ceratobranchial 8 Hypobranchial 8 Basibranchial 8 Pharyngobranchial 6 Urohyal 1

Pelvic Girdle (n=4) Pelvis 2 Pelvic Spines 2 Pectoral Girdle (n=12) Cleithrum 2 Coracoid 2 Postcleithrum 2 Scapula 2 Supracleithrum 2 Posttemporal 2 Vertebrae (n=24) Atlas 1 Thoracic 3 Precaudal 6 Caudal 12 Penultimate 1 Ultimate 1 Fin (n=1) Total pterygiophgore 1st anal pterygiophore 1

435

APPENDIX- 4 Polynemidae skeletal elements in a complete fish.

APPENDIX-4 cont'd

Epibranchial 6 Ceratobranchial 8 Hypobranchial 8 Basibranchial 8 Pharyngobranchial 6 Urohyal 1

Pelvic Girdle (n=4) Pelvis 2 Pelvic Spines 2 Pectoral Girdle (n=12) Cleithrum 2 Coracoid 2 Postcleithrum 2 Scapula 2 Post-temporal 2 Supracleithrum 2 Vertebrae (n=24) Atlas 1 Thoracic 3 Precaudal 6 Caudal 12 Penultimate 1 Ultimate 1 Fin (n=1) 1 dorsal spine 1

437

APPENDIX- 5. Sciaenidae skeletal elements in a complete fish.

Anatomic region/ Expected no. Skeletal element-Cranial Olfactory Region (n=17) Ethmoid 1 Vomer 1 Prefrontal 2 Alisphenoid 2 Parasphenoid 1 Total orbital Series 10 Occipital Region (n=4) Basioccipital 1 Exoccipital 2 Supraoccipital 1 Otic Region (n=16) Opisthotic 2 Epiotic 2 Prootic 2 Pterotic 2 Sphenotic 2 Otolith 6 Investing Bones Region (n=6) Parietal 2 Nasal 2 Frontal 2 Lateral region (n=14) Premaxilla 2 Maxilla 2 Dentary 2 Articular&Angular 4 Preopercle 2 Lachrymal 2 Mandibular region (n=8) Palatine 2 Ectopterygoid 2 Metapterygoid 2 Quadrate 2 Hyoid region (n=12) Hyomandibular 2 Symplectic 2 Interhyal 2 Glosshyal 1 Basihyal 1 Epihyal 2

438

APPENDIX-5cont'd

Ceratohyal Hypohyal 2 Anatomic region/ Expected no. Skeletal element- Opercular Series (n=20) Branchiostegal ray 14 Opercular 2 Interopercular 2 Subopercular 2 Branchial Arch (n=41) pharyngeal plate 4 Epibranchial 6 Ceratobranchial 8 Hypobranchial 8 Basibranchial 8 Pharyngobranchial 6 Urohyal 1

Pelvic Girdle (n=4) Pelvis 2 Pelvic Spines 2 Pectoral Girdle (n=12) Cleithrum 2 Coracoid 2 Postcleithrum 2 Scapula 2 Posttemporal 2 Supracleithrum 2 Vertebrae (n=25) Atlas 1 Thoracic 3 Precaudal 6 Caudal 13 Penultimate 1 Ultimate 1 Fin (n=5) Pectoral spine 2 Dorsal spine 1 Anal spine 1

439

APPENDIX- 6 Clupeidae skeletal elements in a complete fish.

Anatomic region/ Expected no. Skeletal element-Cranial Olfactory Region (n=17) Lateral Ethmoid 2 Vomer 1 Prefrontal 2 Supraethmoid 1 Alisphenoid 2 Parasphenoid 1 Total orbital Series 8

Otic Region (n=4) Basioccipital 1 Exoccipital 2 Supraoccipital 1 Otic Region (n=16) Opisthotic 2 Epiotic 2 Prootic 2 Pterotic 2 Sphenotic 2 Otolith 6 Investing Bones Region (n=6) Parietal 2 Nasal 2 Frontal 2 Lateral region (n=14) Premaxilla 2 Maxilla 2 Dentary 2 Articular&Angular 4 Preopercle 2 Lachrymal 2 Mandibular region (n=8) Palatine 2 Ectopterygoid 2 Metapterygoid 2 Quadrate 2 Hyoid region (n=13) Hyomandibular 2 Symplectic 2 Interhyal 2 Epihyal 2 Ceratohyal 2 Hypohyal 2 Basihyal 1 Anatomic region/ Expected no. Skeletal element- Opercular Series (n=16) Branchiostegal ray 10

440

APPENDIX- 6 cont'd 2 Opercular Interopercular 2 Subopercular 2 Branchial Arch (n=33) pharyngeal plate 2 Epibranchial 8 Ceratobranchial 8 Hypobranchial 6 Basibranchial 2 Pharyngobranchial 6 Urohyal 1

Pelvic Girdle (n=4) Pelvis 2 Pelvic Spines 2 Pectoral Girdle (n=12) Cleithrum 2 Coracoid 2 Postcleithrum 2 Scapula 2

Posttemporal 2 Supracleithrum 2 Vertebrae (n=48) Atlas 1 Thoracic 3 Precaudal 10 Caudal 32 Penultimate 1 Ultimate 1 Fin (n=1) Dorsal spine 1

441

APPENDIX- 7 Tetraodontidae skeletal elements in a complete fish

Anatomic region/ Expected no. Skeletal element-Cranial Olfactory Region (n=8) Ethmoid 1 Vomer 1 Prefrontal 2 Supraethmoid 1 Pterisphenoid 2 Parasphenoid 1 Occipital Region (n=4) Basioccipital 1 Exoccipital 2 Supraoccipital 1 Otic Region (n=10) Epiotic 2 Prootic 2 Pterotic 2 Sphenotic 2 Otolith 2 Investing Bones Region (n=2) Frontal 2 Lateral region (n=12) Premaxilla 2 Maxilla 2 Dentary 2 Articular&Angular 4 Preopercle 2 Mandibular region (n=8) Palatine 2 Ectopterygoid 2 Metapterygoid 2 Quadrate 2 Hyoid region (n=10) Hyomandibular 2 Symplectic 2 Epihyal 2 Ceratohyal 2 Hypohyal 2

442

APPENDIX-7 cont'd

Anatomic region/ Expected no. Skeletal element- Opercular Series (n=8) Branchiostegal ray 2 Opercular 2 Interopercular 2 Subopercular 2 Branchial Arch (n=33) Epibranchial 8 Ceratobranchial 10 Hypobranchial 6 Basibranchial 3 Pharyngobranchial 6 Pectoral Girdle (n=12) Cleithrum 2 Coracoid 2 Postcleithrum 2 Scapula 2 Posttemporal 2 Supracleithrum 2 Vertebrae (n=18) Atlas 1 Thoracic 3 Precaudal 4 Caudal 8 Penultimate 1 Ultimate 1

443

APPENDIX- 8 Belonidae skeletal elements in a complete fish.

Anatomic region/ Expected no. Skeletal element-Cranial Olfactory Region (n=17) Lateral Ethmoid 2 Vomer 1 Prefrontal 2 Supraethmoid 1 Alisphenoid 2 Parasphenoid 1 Total orbital Series 8 Otic Region (n=4) Basioccipital 1 Exoccipital 2 Supraoccipital 1 Otic Region (n=16) Opisthotic 2 Epiotic 2 Prootic 2 Pterotic 2 Sphenotic 2 Otolith 6 Investing Bones Region (n=6) Parietal 2 Nasal 2 Frontal 2 Lateral region (n=14) Premaxilla 2 Maxilla 2 Dentary 2 Articular&Angular 4 Preopercle 2 Lachrymal 2 Mandibular region (n=8) Palatine 2 Ectopterygoid 2 Metapterygoid 2 Quadrate 2 Hyoid region (n=13) Hyomandibular 2 Symplectic 2 Interhyal 2 Epihyal 2 Ceratohyal 2 Hypohyal 2 Basihyal 1 Anatomic region/ Expected no. Skeletal element- Opercular Series (n=16) Branchiostegal ray 10

444

APPENDIX-8 cont'd

Opercular Interopercular 2 Subopercular 2 Branchial Arch (n=31) pharyngeal plate 4 Epibranchial 6 Ceratobranchial 8 Hypobranchial 8 Basibranchial 2 Pharyngobranchial 2 Urohyal 1

Pelvic Girdle (n=4) Pelvis 2 Pelvic Spines 2 Pectoral Girdle (n=12) Cleithrum 2 Coracoid 2 Postcleithrum 2 Scapula 2 Posttemporal 2 Supracleithrum 2 Vertebrae (n=75) Atlas 1 Thoracic 4 Precaudal 41 Caudal 27 Penultimate 1 Ultimate 1

445

APPENDIX- 9 Pristigasteridae skeletal elements in a complete fish.

Anatomic region/ Expected no. Skeletal element-Cranial Olfactory Region (n=17) Lateral Ethmoid 2 Vomer 1 Prefrontal 2 Supraethmoid 1 Alisphenoid 2 Parasphenoid 1 Total orbital Series 8 Otic Region (n=4) Basioccipital 1 Exoccipital 2 Supraoccipital 1 Otic Region (n=16) Opisthotic 2 Epiotic 2 Prootic 2 Pterotic 2 Sphenotic 2 Otolith 6 Investing Bones Region (n=6) Parietal 2 Nasal 2 Frontal 2 Lateral region (n=14) Premaxilla 2 Maxilla 2 Dentary 2 Articular&Angular 4 Preopercle 2 Lachrymal 2 Mandibular region (n=8) Palatine 2 Ectopterygoid 2 Metapterygoid 2 Quadrate 2 Hyoid region (n=14) Hyomandibular 2 Symplectic 2 Interhyal 2 Epihyal 2 Ceratohyal 2 Hypohyal 2 Basihyal 2 Anatomic region/ Expected no. Skeletal element- Opercular Series (n=20) Branchiostegal ray 14 Opercular 2 Interopercular 2 Subopercular 2 446

APPENDIX-9 cont'd

Branchial Arch (n=31) pharyngeal plate 4 Epibranchial 6 Ceratobranchial 8 Hypobranchial 8 Basibranchial 2 Pharyngobranchial 2 Urohyal 1

Pelvic Girdle (n=4) Pelvis 2 Pelvic Spines 2 Pectoral Girdle (n=12) Cleithrum 2 Coracoid 2 Postcleithrum 2 Scapula 2 Posttemporal 2 Supracleithrum 2 Vertebrae (n=48) Atlas 1 Thoracic 3 Precaudal 7 Caudal 26 Penultimate 1 Ultimate 1 Fin (n=1) Dorsal spine 1

447