Maya Exploitation of Animal Resources During the Middle Preclassic Period: an Archeozoological Analysis from Pacbitun, Belize

MAYA EXPLOITATION OF ANIMAL RESOURCES DURING THE MIDDLE PRECLASSIC PERIOD: AN ARCHEOZOOLOGICAL ANALYSIS FROM PACBITUN, BELIZE

A Thesis Submitted to the Committee of Graduate Studies in Partial Fulfillment of the Requirements for the Degree of Master of Arts in the Faculty of Arts and Science

TRENT UNIVERSITY

Peterborough, Ontario, Canada

Anthropology M.A. Graduate Program

January 2014

ABSTRACT

Maya Exploitation of Animal Resources during the Middle Preclassic Period: An Archaeozoological Analysis from Pacbitun, Belize

Arianne Boileau

This study examines the foraging strategies of animal resource exploitation during the Middle Preclassic period (900–300 BC) at the ancient Maya site of Pacbitun, Belize.

The faunal remains analyzed in this study were recovered from various domestic structures associated with the production of shell artifacts. Detailed taphonomic analyses have revealed that the Pacbitun faunal remains were particularly affected by weathering and density-mediated attrition. White-tailed deer was the prey most frequently acquired by the Middle Preclassic Maya of Pacbitun, followed by other lower-ranked artiodactyls.

A variety of less profitable prey were sometimes included in the diet breadth. Using the central place forager prey choice model as a framework, the analysis of diet breadth, habitat use, and carcass transport patterns suggests that most animal resources were acquired from terrestrial habitats, at short distances from the site. Complete carcasses of large game appear to have been frequently transported to the site, where they were exploited for their meat and marrow. Comparisons with other Middle Preclassic faunal assemblages indicate significant differences in terms of taxonomic composition, with an emphasis on the procurement of fish and turtles. It is suggested that the Middle Preclassic

Maya adopted foraging strategies focusing on the exploitation of local habitats, with occasional use of exotic resources.

Keywords: Archaeozoology, subsistence, foraging theory, animal resource exploitation, taphonomy, ancient Maya, Middle Preclassic, Pacbitun, Belize.

ACKNOWLEDGEMENTS

The completion of this thesis would not have been possible without the support of many individuals. First and foremost, I would like to thank my advisor Dr. Eugène Morin, who showed support and endless patience every step of the way. His edits and advice have helped me to become a better writer and researcher. I am deeply thankful for all the time and effort he invested in teaching me and cannot thank him enough for the opportunities he has granted me. I would also like to express my gratitude to Dr. Paul

Healy, who kindly shared his love of Maya archaeology with me. His help, advice, and encouragements are deeply appreciated. I am indebted to Dr. Terry Powis for inviting me to join the Pacbitun Regional Archaeological Project (PRAP) and allowing me to examine the faunal material from Pacbitun. He certainly deserves special mention for answering an immeasurable number of emails concerning the excavations at Pacbitun. I also wish to thank Dr. Carolyn Freiwald who graciously accepted to serve as my external examiner.

Comments and suggestions provided by all the members of my committee contributed to improve this thesis.

I would like to thank the Institute of Archaeology (IOA) in Belize, especially Drs.

Jaime Awe and John Morris, for granting permission to export the faunal material analyzed in this study to Canada. My gratitude also goes to the staff of the Department of

Vertebrate Paleontology at the Royal Ontario Museum, particularly Dr. Kevin Seymour and Brian Iwama, for their warm welcome and for providing access to the ROM vertebrate comparative collections. Further thanks go to fellow Maya archaeozoologists,

Norbert Stanchly and Dr. Erin Thornton, who provided useful information on the identification of vertebrate remains from tropical environments. I also greatly appreciated

iii

and learned much from my discussions with them. I extend my gratitude to Dr. Jocelyn

Williams for her assistance in the identification of human remains.

I am deeply grateful to Dr. Gyles Iannone for inviting me to participate in the

Social Archaeology Research Program (SARP) in Belize and for allowing me to collect animal specimens that are now part of the archaeozoological reference collection at Trent

University. Gyles provided me with my first opportunity to excavate at a Maya site and trusted me in the supervision of an excavation unit, which I am thankful for. This first trip to Belize made me discover an incredible country filled with history and beautiful people.

I never looked back at my decision to study Maya archaeology from that moment.

This project would not have been possible without the financial support from a number of sources. My research was funded by a Joseph-Armand Bombardier Canada

Graduate Scholarship from the Social Science and Humanities Research Council

(SSHRC) and a Fond de Recherche Société et Culture du Québec Master’s scholarship. I also wish to acknowledge the financial support of Trent University which allowed me to complete the present study.

Thanks also go to my fellow anthropology graduate students. In particular, I am grateful to Shannen Stronge and Esther Beauregard for reading over and editing my chapters. I also would like to thank all the people who, over the course of the past three years, have become both friends and colleagues: Steven “Morgan” Moodie, Esther

Beauregard, Kendall Hills, Shannen Stronge, Jodi Schmidt, Kat Elaschuk, Amandah Van

Merlin, Dan Savage, Véronique Belisle, Hannah Schmidt, and Kristine Williams. Their support, help, and humour have made my stay in Peterborough an unforgettable experience! I also wish to thank them all for answering an endless flow of questions about proper English grammar and syntax. iv

Last, but not least, I would like to thank my family. Merci à vous tous pour votre amour, vos encouragements et votre support, car sans eux, je n’aurais pu surmonter toutes les épreuves qui ont entravé mon chemin. Maman et Papa, je vous remercie de n’avoir jamais questionné mon désir de devenir archéologue et pour toujours m’avoir poussée à réaliser mes rêves. Cela a fait toute la différence. Philippe, merci d’être le frère dont j’ai besoin et de nourrir mes heures de travail de bonnes suggestions de musique. Grand-maman, merci de toujours me poser les questions les plus pertinentes concernant mes recherches et de m’avoir incluse dans tes prières.

TABLE OF CONTENTS

Abstract ...... ii Acknowledgements ...... iii Table of Contents ...... vi List of Figures ...... ix List of Tables ...... x

CHAPTER 1 : Introduction ...... 1 1.1 Research objectives ...... 2 1.2 The setting ...... 3 1.3 The Middle Preclassic period ...... 5 1.4 Thesis overview ...... 8

CHAPTER 2 : Middle Preclassic Maya Diet ...... 9 2.1 History of Maya archaeozoology ...... 9 2.2 Ancient Maya diet ...... 11 2.2.1 Plant remains ...... 11 2.2.2 Animal resources ...... 14 2.2.3 Middle Preclassic use of animals ...... 22 2.2.4 Stable isotope analysis ...... 27 2.3 Summary ...... 29

CHAPTER 3 : Site Description and Previous Research ...... 30 3.1 Geographical context ...... 30 3.2 Archaeological investigations ...... 35 3.3 Site chronology ...... 38 3.4 Middle Preclassic investigations ...... 41 3.5 Artifactual assemblages from the Middle Preclassic period ...... 46 3.6 Subsistence practices ...... 48 3.7 The Middle Preclassic at Pacbitun: A summary ...... 51

CHAPTER 4 : Foraging Theory ...... 53 4.1 Theoretical approach ...... 53

4.1.1 Prey choice model ...... 54 4.1.2 Patch choice models ...... 55 4.1.3 Central place forager prey choice model ...... 57 4.2 Archaeological applications of foraging theory ...... 59 4.2.1 Use of foraging models in this study ...... 59 4.2.2 Prey rankings ...... 60 4.2.3 Abundance indices ...... 64 4.3 Animal ecology and behavior ...... 65

CHAPTER 5 : Methodology ...... 75 5.1 Definitions and identification procedures ...... 75 5.2 Quantification methods ...... 76 5.2.1 Strengths and weaknesses of NISP, MNE, and MNI ...... 78 5.2.2 Use of quantification methods in this study ...... 81 5.3 Refitting ...... 82 5.4 Age and sex ...... 83 5.5 Taphonomic modifications ...... 85 5.5.1 Fractures ...... 85 5.5.2 Butchery and tool use ...... 86 5.5.3 Carnivore ravaging ...... 88 5.5.4 Fragmentation ...... 90 5.5.5 Burning ...... 91 5.5.6 Additional taphonomic agents ...... 92 5.6 Summary ...... 94

CHAPTER 6 : Sample Description and Taphonomy ...... 95 6.1 The Pacbitun faunal assemblages ...... 95 6.2 Taphonomy ...... 99 6.2.1 Testing the stratigraphic sequence ...... 100 6.2.2 Recovery methods ...... 102 6.2.3 Density-mediated attrition ...... 104 6.2.4 Bone burning ...... 108 6.2.5 Post-depositional destruction ...... 109 6.2.6 Bone surface preservation ...... 111 6.2.7 Human and carnivore agents ...... 114

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6.2.8 Taphonomy of fish and birds ...... 119 6.3 Summary ...... 121

CHAPTER 7 : Results ...... 123 7.1 Taxonomic composition ...... 123 7.2 Skeletal part representation ...... 125 7.3 Mortality profiles ...... 129 7.4 Scheduling of activities ...... 133 7.5 Diet breadth ...... 133 7.6 Habitat use ...... 139 7.7 Transport selectivity ...... 142 7.8 Processing of skeletal parts ...... 147 7.9 Pacbitun foraging strategies: A discussion ...... 149 7.10 Subsistence strategies in the southern Maya lowlands ...... 152

CHAPTER 8 : Conclusion ...... 160 8.1 Research summary ...... 160 8.2 Limitations and significance ...... 162 8.3 Future Directions ...... 163

Bibliography ...... 165 Appendix A ...... 200

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LIST OF FIGURES

Figure 1.1 Map of the Maya subarea including sites discussed in the present study...... 4

Figure 3.1 Map of the Belize Valley ...... 31

Figure 3.2 Location of Pacbitun in the Upper Belize Valley ...... 32

Figure 3.3 Map of the Pacbitun settlement survey ...... 34

Figure 3.4 Plan of the epicenter of Pacbitun...... 35

Figure 3.5 Core zone of Pacbitun ...... 36

Figure 3.6 Chronology and ceramic complexes of Pacbitun in comparison to those of Barton Ramie, Xunantunich, and Cahal Pech...... 39

Figure 3.7 Plan of the Middle Preclassic sub-structures in Plaza B ...... 43

Figure 3.8 Plan of the Middle Preclassic Sub-Structure A-1 in Plaza A...... 45

Figure 4.1 Ranking of mammals at Pacbitun according to body mass ...... 63

Figure 6.1 Fragment size distribution by screen size for all faunal specimens in the Pacbitun Middle Preclassic assemblages...... 103

Figure 6.2 %NNISP of long bone portions of white-tailed deer versus bone density values (g/cm3) ...... 107

Figure 6.3 %MAU of long bone portions of white-tailed deer versus bone density values (g/cm3) ...... 107

Figure 6.4 Gnaw and cut marks on the distal end of a right white-tailed deer femur ..... 118

Figure 7.1 White-tailed deer body part representation in the Middle Preclassic assemblages at Pacbitun ...... 126

Figure 7.2 Comparison of white-tailed deer element frequencies (%NNISP) in the Pacbitun assemblages with the reindeer MUI, grease index, and UMI...... 145

LIST OF TABLES

Table 2.1 Plant remains recovered from Middle Preclassic deposits at southern lowland Maya sites...... 12

Table 2.2 List of identified taxa recovered from Middle Preclassic deposits at southern lowland Maya sites (NISP counts)...... 23

Table 3.1 Possible source location for local and exotic raw materials found at Pacbitun.47

Table 3.2 Vertebrate and invertebrate remains identified by Stanchly (1999) ...... 49

Table 4.1 Data and references for body mass of mammalian taxa at Pacbitun...... 63

Table 5.1 Taxonomic groups based on body size...... 76

Table 6.1 Number of specimens by primary and secondary contexts for the Middle Preclassic assemblages...... 96

Table 6.2 Pre- and post-refit NISP counts by time period...... 96

Table 6.3 Distribution of faunal remains by zoological class for the early and late Middle Preclassic samples at Pacbitun...... 97

Table 6.4 Identified taxa by NISP and MNI for the early and late Middle Preclassic samples at Pacbitun...... 98

Table 6.5 Taxonomic representation in the Middle Preclassic samples by mesh size, in percentages...... 104

Table 6.6 Bone density values of Rangifer tarandus compared to %NNISP and %MAU values for white-tailed deer long bone portions in the Middle Preclassic Pacbitun assemblages...... 106

Table 6.7 Degree of post-depositional completeness by time period...... 110

Table 6.8 NISP counts for green- and dry-bone fractures by time period...... 111

Table 6.9 Overall surface state for the Middle Preclassic Pacbitun assemblages...... 112

Table 6.10 Percentage of observable surface in the Middle Preclassic samples...... 113

Table 6.11 Frequencies of cutmarks on identified specimens and indeterminate long bone shafts by overall surface state for the Middle Preclassic samples at Pacbitun...... 114

Table 6.12 Frequencies of anthropogenic marks observed in the Middle Preclassic Pacbitun assemblages...... 115 x

Table 6.13 Frequencies of carnivores marks observed in the Middle Preclassic Pacbitun assemblages...... 117

Table 6.14 Extent of carnivore gnawing on bone surfaces for the Middle Preclassic samples at Pacbitun...... 118

Table 7.1 Skeletal part frequencies for white-tailed deer in the Middle Preclassic assemblages from Pacbitun ...... 127

Table 7.2 Skeletal part frequencies (NISP) of armadillo, peccary, and red brocket deer for the Middle Preclassic period at Pacbitun...... 128

Table 7.3 Number of specimens identified per category of epiphyseal fusion for white- tailed deer in the early and late Middle Preclassic samples at Pacbitun ...... 131

Table 7.4 Habitat fidelity values for the mammalian species identified in the Middle Preclassic assemblages at Pacbitun ...... 140

Table 7.5 Analysis of habitat fidelity for the Pacbitun assemblages, by time period ..... 140

Table 7.6 Spearman’s rank order correlations between skeletal part representation and the MUI, grease index, and UMI...... 146

Table 7.7 Percentages of vertebrate taxa identified in Middle Preclassic assemblages at Pacbitun, Tolok group at Cahal Pech, Colha and Bayak ...... 154

Table 7.8 Results of the Kolmogorov-Smirnov tests for the Middle Preclassic assemblages from the southern lowlands...... 156

Table 7.9 Abundances of marine fish at southern lowlands Maya sites during the Middle Preclassic period, with average distances from the coast of the Caribbean Sea ...... 158

CHAPTER 1: INTRODUCTION

For decades, archaeological research on the ancient Maya focused almost exclusively on documenting the ways of life of the Classic period (AD 300–900).

Although these efforts contributed significantly to our understanding of the Maya, little was known about the early developments of this civilization during the Preclassic period

(2000 BC–AD 250). This situation was also partly attributable to the Mesoamerican tradition of building new structures on top of older ones, which made it difficult for archaeologists to access Preclassic cultural remains, as they were often buried under meters of construction fill. However, in the past thirty years, more research projects have focused on documenting the social, political, and economic systems of the Preclassic

Maya at sites such as Blackman Eddy (Garber et al. 2004a; Garber et al. 2004b), Cahal

Pech (Healy and Awe 1995b, 1996; Powis et al. 1999; Healy et al. 2004), Cuello

(Hammond 1991, 2005), Colha (Hester et al. 1982, 1994), and Cival (Estrada-Belli 2011).

Unfortunately, growing interest in the excavations of Preclassic structures did not go in tandem with increased attention aimed at the recovery of faunal remains. As a result, the strategies of animal resource exploitation of the Preclassic Maya remain poorly documented. Several research projects have produced detailed analyses of animal procurement strategies for sites located in northern Belize (e.g., Shaw 1991; Masson

2004b; Carr and Fradkin 2008), but information for inland sites located in the Belize

River Valley or the Peten region of Guatemala remains scarce. This problem is also exacerbated by the small sample size of many faunal assemblages, which often precludes detailed investigation of animal use during this early time period.

1.1 Research objectives

The goal of this study is to contribute to filling this gap in research by investigating the strategies of animal resource exploitation adopted by the Preclassic

Maya in the southern lowlands. This issue is examined using faunal remains recovered from Middle Preclassic (900–300 BC) deposits at Pacbitun, an ancient Maya site located at the southern rim of the Belize River Valley and occupied from the Middle Preclassic to

Terminal Classic periods. Faunal remains recovered during the 1995 and 1996 field seasons have previously been presented in a preliminary report by Norbert Stanchly

(1999). The present study includes the material from the 1997 and 2008–2011 excavations to provide a fuller archaeozoological analysis of the Middle Preclassic faunal remains from Pacbitun. The objectives of this analysis are as follows:

1) To complete a taphonomic analysis of the faunal remains and evaluate the impact

of anthropogenic and natural agents on the assemblages;

2) To characterize the foraging strategies of the Maya of Pacbitun during the Middle

Preclassic period through an investigation of diet breadth, habitat use, transport

decisions, and skeletal part processing;

3) To consider how strategies of animal resource exploitation at Pacbitun contrast

with that of other Middle Preclassic sites in the southern Maya lowlands.

This study will examine the relationship that existed between the ancient Maya and the animal populations of the Belize River Valley during the Middle Preclassic.

Additionally, this research will enhance our overall understanding of how the early Maya of the southern lowlands interacted with their environment. Although this research relies strictly on empirical faunal data to examine foraging strategies, the author would like to

3 remind that cultural beliefs about animals also play an important role in food choices and overall animal use. However, these considerations are outside the scope of this study. It should also be noted that the data presented in this study focus exclusively on the southern lowlands. Although more attention has recently been directed at documenting

Middle Preclassic occupations in the northern lowlands (e.g., Rissolo et al. 2005; Stanton and Ardren 2005; Anderson 2011), information on the exploitation of animals resources in this region remains scanty. In fact, the only published faunal analysis of vertebrate remains known to the author from this region is that of Dzibilchaltun by Wing (1980).

The sample for this site was considered too small (NISP = 53) to be included in this study. The next sections of this chapter provide general information on the Maya subarea and the cultural developments of the Middle Preclassic period.

1.2 The setting

The ancient Maya inhabited the southeastern portion of Mesoamerica for over two millennia (2000 BC–AD 1500). During this period, the Maya subarea (Figure 1.1) encompassed southeastern Mexico, Guatemala, Belize, and the western regions of

Honduras and El Salvador (Demarest 2004:11). The subarea is generally divided into three loosely defined environmental regions: the highlands, northern lowlands, and southern lowlands (Sharer 1994:20; Demarest 2004:121).

The Maya highlands are formed by the mountainous region which stretches from the highlands of Chiapas, Mexico, to El Salvador. This region is characterized by valleys and basins of rich and fertile soils which lie above 800 m in elevation (Sharer 1994:26–

32; Demarest 2004:121). The northern lowlands are formed by the northern section of the

Yucatan peninsula of Mexico and comprise the states of Campeche, Quintana Roo, and

Yucatan. The terrain is relatively flat, with the exception of the Puuc Hills. The region is characterized by a dry environment; there is little rainfall (500–2000 mm) and surface

Figure 1.1 Map of the Maya subarea including sites discussed in the present study. Map modified from Brown and Witschey (2008), retrieved from: http://mayagis.smv.org/maps_of_the_maya_area.htm.

5 water is generally scarce, with the exception of natural sinkholes. As a result, scrub vegetation prevails and soils are generally thin and show low fertility (Demarest

2004:120–127; McKillop 2004a:29–34).

The southern lowlands—the focus of this study—separate the highlands from the northern lowlands and include the country of Belize, the Peten district of Guatemala, parts of Chiapas and lowland Honduras (McKillop 2004a:29). This region presents a more hilly terrain, with the Maya Mountains constituting the highest elevation on the landscape. The southern lowlands receive abundant rainfall (2000–3000 mm) and, as a result, are covered with lush rainforests. Large rivers and their tributaries also cover the landscape. These would have provided access to potable water and may have constituted important navigation routes in ancient times. Soils can be quite fertile in this region, especially in floodplains (Sharer 1994:34; Demarest 2004:120–127; McKillop 2004a:29–

34).

1.3 The Middle Preclassic period

Despite its importance for understanding the origins of the Maya civilization, the

Preclassic period (2000 BC–AD 250) is poorly documented in comparison to the Classic

(AD 250–900) and Postclassic (AD 900–1525) periods (Powis 2005; Healy 2006).

Archaeological evidence from the Early Preclassic (2000–1000 BC) indicates that a handful of farming villages were established in the southern lowlands at sites such as

Cahal Pech, Nakbe, Seibal, Cival, Tikal, and Blackman Eddy (Figure 1.1). The architectural remains identified at these sites are modest (Estrada-Belli 2011). Domestic pole-and-thatch structures were generally erected directly on, or slightly above, the ground surface (Hansen 1998; Garber et al. 2004a; Healy et al. 2004). It is believed that

6 little social or economic differentiation existed within these communities, although larger villages might have been headed by a chief (Garber et al. 2004a; Healy et al. 2004).

The farming village lifeway established during the Early Preclassic likely continued during the Middle Preclassic period (1000–300 BC). This latter period, however, was marked by the first indications of increased social and economic complexity, even though these signs are subtle at most sites (Healy 2006). In fact, the

Middle Preclassic period may have coincided with a transition from a relatively egalitarian to a ranked society (Healy 2006). Most scholars divide the Middle Preclassic into two sub-periods: the early Middle Preclassic (1000–600 BC) and the late Middle

Preclassic (600–300 BC).

During the early Middle Preclassic, increasing complexity can be found in architecture, as wooden structures were being built on top of platforms made of modest stone walls and covered with plastered floors. These constructions were commonly organized around an open plastered-surface patio. It is believed that these clusters of structures functioned as residential groups, perhaps for an extended family (Hammond and Gerhardt 1990). Examples of these structures were found at Blackman Eddy (Garber et al. 2004a), Cahal Pech (Healy et al. 2004), Cuello (Hammond and Gerhardt 1990), and

Nakbe (Hansen 1998), among others. The emergence of ritual ideology is possibly evidenced by the presence of dedication and termination caches in some of these structures (Garber et al. 2004a; Estrada-Belli 2011). Increasingly elaborate architecture also appears in the form of E-Groups at Seibal (Inomata et al. 2013), Tikal (LaPorte and

Fialko 1995), and Cival (Estrada-Belli 2011). Considered as the first form of public architecture in the lowlands, the construction of these large structures demonstrates the ability of an emerging elite to marshal increasing amounts of labor (Estrada-Belli 2011).

It is hypothesized that E-Groups may have been constructed as a way to legitimize and express the political authority of new elite groups on the landscape (Doyle 2012).

Signs of social and economic complexity became more apparent during the late

Middle Preclassic, a period that witnessed the development of monumental public architecture at several sites, including Cahal Pech (Healy et al. 2004), Itzan (Johnston

2006), and Cuello (Hammond 1991, 2005). Large blocks of cut stones were used for the first time in the lowlands at Nakbe (Hansen 1998), whereas the first sculpted architectural decoration appeared in the form of stucco masks at Blackman Eddy (Garber et al. 2004a).

These larger structures were some of the first occurrences of civic-ceremonial architecture and possibly served as places for public ritual and performance (Healy 2006).

Trade networks within and outside the Maya subarea also intensified during the Middle

Preclassic period. Exotic goods, such as greenstone, obsidian, and marine shells, were obtained from the Motagua Valley, Guatemalan highlands, and Caribbean coast (Garber et al. 2004a; Healy et al. 2004; Hammond 2005; Healy 2006). The early occurrence and diverse origins of these goods suggest the existence of extensive systems of long distance trade and exchange by the beginning of the first millennia BC.

It should be noted that not all Middle Preclassic sites developed at the same rate.

Indeed, many of the cultural developments mentioned above for the early Middle

Preclassic only took place during the late Middle Preclassic or even the Late Preclassic periods at a majority of sites (e.g., Buenavista del Cayo, Ball and Taschek 2004). Overall, the religious, artistic, and architectural developments of the Middle Preclassic period appear to have provided a strong foundation for the political and cultural changes observed during the Late Preclassic (300 BC–AD 250). It is during this time period that we see throughout the Maya lowlands the first recognizable depictions of Maya kings and

8 the establishment of hereditary dynasties that would characterize the Maya civilization for centuries (Healy 2006; Estrada-Belli 2011).

1.4 Thesis overview

This thesis is organized in eight chapters covering different aspects of my research. Chapter 2 offers a general overview of Maya subsistence strategies during the

Middle Preclassic period. Chapter 3 introduces the site of Pacbitun and discusses the research conducted at the site. Chapter 4 reviews the foraging models used to interpret the faunal remains and provides a summary of the ecology and behavior of the prevalent animal species identified at the site. The methods used to quantify the faunal assemblages and assess the integrity of the samples are presented in Chapter 5, whereas Chapter 6 describes the samples analyzed in this study and considers the taphonomic history of the faunal remains. Chapter 7 presents the results of this study and discusses their implications for our understanding of animal resource exploitation at Pacbitun during the

Middle Preclassic period. This chapter also considers how the faunal data align with research conducted at other Middle Preclassic sites in the southern lowlands. Lastly,

Chapter 8 discusses the significance of the results and offers suggestions for future research.

CHAPTER 2: MIDDLE PRECLASSIC MAYA DIET

This chapter provides a comprehensive account of previous research regarding

Maya subsistence during the early developments of this civilization. It begins with a summary of Maya archaeozoology. The second half of this chapter presents the results of botanical, faunal, and isotopic studies relevant to the Maya diet during the Middle

Preclassic period (1000–300 BC).

2.1 History of Maya archaeozoology

The first archaeozoological studies in the Maya area were performed in the 1930s, when the Carnegie Institution of Washington and the Museum of Zoology of the

University of Michigan jointly funded archaeological and biological research in British

Honduras (modern-day Belize) and Guatemala (Chase et al. 2004; Emery 2004c, 2010).

Alongside archaeological fieldwork, the emphasis was put on collecting biological specimens of vertebrate and invertebrate faunas of the Neotropics (Hubbs 1935; Murie

1935; Stuart 1935; van Tyne 1935; Goodrich and van der Schalie 1937) .

The importance of animal resources for the ancient Maya was first discussed in excavation reports from the sites of Uaxactun (Ricketson 1937), Piedras Negras (Coe

1959), and Holmul (Merwin and Vaillant 1932). Early faunal studies were generally carried out by zoologists who limited their analysis to the taxonomic identification of specimens, producing species lists that were appended to site reports (Kidder et al.

1946:152–157; Moedano-Koer 1946; Woodbury and Trik 1954; Pina-Chan 1968).

Typically, only specimens recovered from special deposits and artifacts made of bone and shell were analyzed. The results of the faunal studies were rarely integrated into interpretative frameworks (Emery 2004c, 2010). Although a considerable amount of

10 research was devoted to studying Maya subsistence strategies in the 1950s and 1960s, archaeozoological studies remained fairly limited in scope. Because it was largely assumed that animal proteins did not play an important role in the Maya diet, the study of animal resources was not considered an important avenue of research (Clutton-Brock and

Hammond 1994; Emery 2004c, 2010).

Under the influence of the “New Archaeology,” the analysis of faunal remains was progressively integrated into discussions regarding the paleoenvironment and subsistence of the ancient Maya (e.g., Savage 1971; Olsen 1972; Andrews et al. 1974;

Luther 1974; Wing 1975; Pohl 1976; Olsen 1978; Hamblin 1984). Simultaneously, Maya archaeozoologists developed strategies to mitigate problems of sample recovery, preservation, and quantification (Clutton-Brock and Hammond 1994; Emery 2004c). As a result of these concerns, archaeologists began using finer mesh sieving (e.g., 1/8 and 1/16 inch mesh screen), which facilitated the recovery of fish and molluscs specimens from archaeological contexts. This, in turn, stimulated research on the importance of marine and riverine resources to the Maya diet (Moholy-Nagy 1963; Andrews 1969; Lange 1971;

Moholy-Nagy 1978; McKillop 1984; Hamblin 1985; McKillop 1985; Moholy-Nagy

1985; Healy et al. 1990).

Since the 1980s, archaeozoological studies form an integral component of most

Maya archaeological projects. In addition to the study of subsistence and dietary adaptations to environmental changes (Carr 1985, 1986; Cliff and Crane 1989; Pohl 1990;

Wing and Scudder 1991; Powis et al. 1999; Shaw 1999; Teeter 2001; Carr and Fradkin

2008; Götz 2008; Emery 2010), several new themes have emerged, such as the investigation of social structure and differential access to animal resources (Pohl 1994;

Emery 1999; Shaw 1999; Teeter 2001; Collins 2002; Emery 2003; Götz 2009). The use

11 of animals in rituals and feasting has become a popular topic of study (Pohl 1981, 1983;

Carr 1985; Pohl 1990; Masson 1999; Teeter 2001; Montero-Lopez 2009). Transport and exchange of animal resources are also investigated (Carr 1996; Emery 1999; Masson and

Lope 2008; Thornton 2011), as well as the origins and process of animal domestication and husbandry (Hamblin 1984; Pohl 1990; Clutton-Brock and Hammond 1994; Carr

1996; Masson and Lope 2008). Isotopic analyses of animal remains have recently become an additional component of archaeozoological studies (Emery et al. 2000; White et al.

2001b; White 2004; Emery and Thornton 2008a; Repoussard 2009; Freiwald 2010;

Thornton 2011).

2.2 Ancient Maya diet

Information about the subsistence of the Maya during the Middle Preclassic period

(1000–300 BC) is quite limited. This partly results from research biases, as archaeological projects in the study area have largely focused on the development of major centres during the Classic period (AD 300–900) (Healy and Awe 1995a; Healy 1999a; Powis

2005; Healy 2006). Additionally, organic remains are often scarce, perhaps due to unfavorable conditions of preservation. The following sections summarize our understanding of the Maya diet during the Middle Preclassic period by considering botanical, faunal, and isotopic data recovered from sites located in the southern lowlands.

2.2.1 Plant remains

Ethnohistorical accounts (e.g., Tozzer 1941) indicate that the Maya diet was traditionally based on the exploitation of three cultigens: maize (Zea mays), beans

(Phaseolus vulgaris), and squash (Cucurbita spp.). Together, they formed the

Mesoamerican triumvirate (Turner and Miksicek 1984; Lentz 1999; Tykot 2002; Lentz et

12 al. 2005). Research conducted in the Maya subarea (Figure 1.1) suggests that maize and squash were domesticated around 3500 BC and 1500 BC, respectively (Pohl et al. 1996).

The introduction of the common bean appears to date to the Middle Preclassic, although its utilization may be older (Lentz 1999; Lentz et al. 2005). The recovery of beans from the archaeological record is relatively sparse, possibly because they do not preserve well in the humid tropics (Lentz 1999). All three taxa have been recovered in Middle

Preclassic deposits at southern lowland Maya sites (Table 2.1).

Table 2.1 Plant remains recovered from Middle Preclassic deposits at southern lowland Maya sites.

Cahal Pulltrouser Taxon Colha Copan Cuello Tikal Pech Swamp Triumvirate Maize x x x x x x Beans x x Squash x x x x x Root crops Manioc x x x Malanga x Other plants Avocado x x x x Bottle gourd x Cacao x x x Cashew x Chili peppers x x Coyol palm x x Fig x Guava x x x Hogplum x x x Kinep x Mamey x x x Nance x x Ramón x Source: Cahal Pech (Lawlor et al. 1995; Powis et al. 1999; Wiesen and Lentz 1999; Lentz et al. 2005), Colha (Turner and Miksicek 1984; Jones 1994), Copan (Turner and Miksicek 1984; Lentz 1991), Cuello (Miksicek et al. 1981a; Miksicek 1991), Pulltrouser Swamp (Turner and Miksicek 1984; Pohl et al. 1996), Tikal (Turner and Miksicek 1984).

In addition to the triumvirate, the Maya relied on tree cropping (Puleston 1982;

Turner and Miksicek 1984; Lentz 1991), as is attested by the recovery of archaeobotanical remains of domesticated fruit trees, such as the avocado (Persea americana), cashew (Anacardium occidentale), and cacao (Theobroma cacao) (Lentz

1999) (Table 2.1). A variety of fruits, including the fig (Ficus spp.), guava (Psidium guajava), hogplum (Spondias mombin), kinep (Talisa oliviformis), mamey (Calocarpum mammosum), and nance (Byrsonima crassifolia), as well as chili peppers (Capsicum annuum), were likely exploited for food (Roys 1931:346–348; Turner and Miksicek 1984;

Miksicek 1991; Lentz 1999; Powis et al. 1999; Wiesen and Lentz 1999; Colunga-García

Marín and Zizumbo-Villarreal 2004; Lentz et al. 2005). These were likely cultivated in orchards and gardens (Turner and Miksicek 1984; Lentz 1991, 1999) or gathered in the forest (Lentz 1999).

Although Bronson (1966) has argued that root crops might have been important staples in Maya agriculture, the degree to which they contributed to the Maya diet is still debated (Lentz 1999). Sweet potato (Ipomoea batatas), yam bean (Pachyrhizus erosus), malanga (Xanthosoma sp.), and manioc (Manihot esculenta) are frequently mentioned in ethnohistoric records (e.g., Roys 1931:346; Tozzer 1941:196). However, they are rarely recovered from archaeological sites, again likely due to poor preservation (Turner and

Miksicek 1984; Hather and Hammond 1994; Lentz 1999). Certain plants, such as the coyol (Acrocomia aculeata), cohune (Attalea cohune), and ramón (Brosimum alicastrum), were possibly consumed during times of famine (Tozzer 1941:200; Miksicek et al. 1981b;

Marcus 1982; Turner and Miksicek 1984; Lentz 1991; McKillop 1996; Lentz 1999;

Wiesen and Lentz 1999). In addition to being exploited for food, many of these plants

14 may have been used as medicine, animal fodder, and construction material (Roys 1931;

Tozzer 1941).

2.2.2 Animal resources

Animal species diversity in the Maya subarea is one of the richest in the world

(Emmons 1997; Emery 2010:45). Archaeological, ethnohistoric, and iconographic evidence indicates that animals had both socioeconomic and symbolic importance for the residents of this region from the Late Preclassic to Colonial periods (Tozzer 1941; Pohl

1983; Hamblin 1984; Emery 2010). However, during the Middle Preclassic, animals seem to have been used primarily for food and utilitarian purposes (Willey 1978; Pohl 1990;

Shaw 1991; Wing and Scudder 1991; Stanchly 1995; Moholy-Nagy 1998; Powis et al.

1999; Harrigan 2004; Masson 2004a, 2004b; Carr and Fradkin 2008; Emery 2010). It is probable that the Middle Preclassic Maya made use of animals for religious purposes, but little archaeological data is currently available to support this idea. One of the only examples come from Cahal Pech, where several hundred complete specimens of apple snails were discovered in a cache (Stanchly 1995). At Blackman Eddy, a concentration of faunal remains associated with multiple serving vessels was interpreted as the remnants of a feasting event (Brown 2007). Therefore, in order to present a complete range of the possible roles animals may have played in the Middle Preclassic society, this section presents the main taxa exploited by the ancient Maya, including information on the use of animals in rituals from later time periods.

Artiodactyls were among the most valuable animal resources used by the ancient

Maya. The white-tailed deer (Odocoileus virginianus)1, red brocket deer (Mazama americana), collared peccary (Pecari tajacu), and white-lipped peccary (Tayassu pecari) are the most common taxa recovered in Middle Preclassic deposits (Pohl 1990; Shaw

1991; Wing and Scudder 1991; Stanchly and Dale 1992; Stanchly 1995; Powis et al.

1999; Carr and Fradkin 2008; Emery 2010). They seem to have been a favoured source of food and raw material (Tozzer 1941:204; Pohl 1990; Hopkins 1992; Carr 1996; Teeter

2001:215–217; Emery 2010). Specifically, the white-tailed deer is regarded as the main animal staple in the Maya diet (Pohl 1990; Emery 2010). Commonly used in sacrifices during the Classic and Postclassic periods, deer legs and heads, as well as peccary heads, are depicted as offerings in the Maya codices2 (Tozzer and Allen 1910; Tozzer 1941:115,

163–165; Pohl 1983). Deer skulls were also recovered in ritual caches dating to the Late

Preclassic at Cuello (Wing and Scudder 1991). Ethnohistoric accounts suggest that the

Postclassic Maya managed deer and peccary herds (Tozzer 1941:127; Hamblin

1984:132–133; Donkin 1985; Carr 1996). It is believed that this activity involved feeding the animals with significant quantities of maize (Emery et al. 2000; van der Merwe et al.

2000). However, stable isotope analyses of carbon and nitrogen on deer and peccary bones do not support this hypothesis for the Middle Preclassic period, as both species exhibit a diet composed predominantly of wild plants (Tykot et al. 1996; Emery et al.

2000; van der Merwe et al. 2000; White et al. 2001b).

1 Latest taxonomic nomenclature retrieved from the Integrated Taxonomic Information System (www.itis.gov, accessed March 28, 2013). 2 The Maya codices are screen-fold books written by the ancient Maya during the Late Postclassic period (AD 1250–1520). The books are made of bark paper on which glyphs and pictures were painted. They record ritual and astronomical information (Vail and Aveni 2004).

Dogs (Canis lupus familiaris) seem to have been the only domesticated animal sued by the Maya during the Middle Preclassic, likely serving both as pets and hunting companions (Tozzer 1941:203; Hamblin 1984; Pohl 1990; White et al. 2001b).

Associated with death and the journey to the underworld, dogs played a large role in

Maya religion during the Classic and Postclassic periods. For instance, they were frequently offered as sacrificial victims in rites associated with annual renewal ceremonies (Tozzer and Allen 1910; Tozzer 1941:143; Pohl and Feldman 1982; Pohl

1983; Hamblin 1984:117–120; White et al. 2001b). Dog skulls and teeth were also recovered as offerings in burial contexts dating to the Late Preclassic (Carr 1986:254;

Valdez 1995). In a society where hunting was the primary means of acquiring proteins and fats, domesticated dogs possibly constituted a reliable source of food when other resources were scarce (Tozzer 1941:203; Pohl 1990; Clutton-Brock and Hammond 1994;

White et al. 2001b). Stable isotope analyses conducted on dog remains recovered from middens dating to the Middle Preclassic period showed large variations in results, suggesting that dogs were not fed a stable household diet, but more likely scavenged, hunted, and foraged on their own (Tykot et al. 1996; van der Merwe et al. 2000; White et al. 2001b). Dogs recovered from special deposits, however, contrast with other dogs as their diet was rich in maize. It was suggested that these animals may have been purposefully fattened for sacrificial purposes (Tykot et al. 1996; White et al. 2001b).

Another canid, the gray fox (Urocyon cinereoargenteus), is also present in the Maya subarea and might have been hunted for its pelt (Teeter 2001) and for food (Hamblin

1984:151–152).

Some small game species also seem to have been economically important for the

Maya during the Middle Preclassic (Pohl 1990; Shaw 1991; Wing and Scudder 1991;

Stanchly and Dale 1992; Stanchly 1995; Powis et al. 1999; Masson 2004a; Carr and

Fradkin 2008; Emery 2010). The armadillo (Dasypus novemcinctus) was probably hunted for its meat as well as for its carapace, which might have been used as a container or instrument (Tozzer 1941:204; Hopkins 1992; Emery 2010). Rabbits, including the cottontail (Sylvilagus floridanus) and forest rabbit (S. brasiliensis), may have been exploited for their meat and pelt (Tozzer 1941:204; Hopkins 1992; Emery 2010). Two large rodents, the agouti (Dasyprocta punctata) and paca (Cuniculus paca), were also probably hunted for their fatty meat (Stuart 1964; Hopkins 1992; Emery 2010).

The use of other mammals as a source of food is speculative. Only identified at a handful of Middle Preclassic sites (see Wing and Scudder 1991; Shaw 1999; Carr and

Fradkin 2008), it is uncertain if the pocket gopher (Orthogeomys hispidus) was considered as a food source or if it is intrusive to the archaeological record (Hopkins

1992; Emery 2010). Other rodents, such as rats and mice, are more likely to have entered archaeological deposits on their own, given that they often live in burrows (Pohl 1983;

Emery 2010). Pohl (1983), however, suggests that they might have held some symbolic importance for the Maya because of their occasional occurrence in ceremonial deposits.

Ethnohistoric accounts indicate that the common opossum (Didelphis marsupialis) and

Virginia opossum (D. virginiana) were used as food sources (Hamblin 1984:154; Teeter

2001; Emery 2010). Similarly, Bishop de Landa reports that the Maya occasionally hunted tapirs (Tozzer 1941:203). Procyonids, including raccoons (Procyon lotor), kinkajous (Potos flavus) and coatis (Nasua narica), are frequent agricultural pests today and, perhaps for this reason, were often used to depict human thieves and gluttons in the

Maya codices (Tozzer 1941:205; Emery 2010). Landa reports that coatis were raised by

Maya women as pets and were sometimes eaten (Tozzer 1941:204–205; Hamblin

1984:147–149). Although all these animals seem to have been exploited during the

Postclassic and Colonial periods, their use during the Middle Preclassic is unclear.

Felines, including the puma (Puma concolor), jaguarundi (Puma yagouaroundi), margay (Leopardus wiedii), and ocelot (Leopardus pardalis), played an essential role in

Maya religion and were often sacrificed in ceremonies during the Classic and Postclassic periods (Tozzer 1941:163; Hopkins 1992). The jaguar (Panthera onca), in particular, has long been associated with sorcery and the supernatural underworld (Tozzer and Allen

1910:355–358; Pohl 1983; Hamblin 1984:164; Emery 2010). Jaguar skins and teeth were important status markers during the Classic period. Priests and rulers appear to have worn them as a sign of their divine power (Pohl 1983; Hopkins 1992; Emery 2010), whereas lesser elites may have used ocelot and margay skins as a display of their status (Emery

2010). It is not known if felines were used in a similar fashion during the Middle

Preclassic period.

Galliforms, including the curassow (Crax rubra), crested guan (Penelope purpurascens), and ocellated turkey (Meleagris ocellata), were probably the most important birds used as food sources (Hamblin 1984; Emery 2010:91). Also possibly hunted for meat were the ground-dwelling tinamous (Tinamidae), ducks and their relatives (Anatidae), herons (Ardeidae), cranes (Gruidae), rails (Rallidae), pigeons and doves (Columbidae) (Tozzer 1941; Shaw 1991:201–203). Although none of these birds, with the exception of the wild turkey (Meleagris gallopavo), were domesticated at the time of conquest, ethnohistoric sources suggest that birds were occasionally captured and penned. The Maya may also have stolen eggs from nests in order to raise the chicks

(Tozzer 1941:202; Hamblin 1984:93–94). Many birds, the turkey above all, held special symbolic importance for the Classic and Postclassic Maya. Parts from these birds were

19 repeatedly used as offerings or deposited in caches and burials (Tozzer and Allen

1910:325–327; Tozzer 1941; Pohl 1983; Hamblin 1984:95; Emery 2010), while colorful feathers were used to decorate clothing, headdresses, and banners (Tozzer 1941:89;

Hamblin 1984:95; Emery 2010). Although they possibly represented a valuable source of food, bird remains are usually rare at Maya sites, perhaps because their fragile bones do not preserve well in tropical environments.

Disagreement generally surrounds the interpretation of amphibians recovered from

Maya sites. Although some pre-Hispanic groups, such as the Campa and Wayapi, seem to have eaten frogs and toads, there is no solid archaeological or ethnohistoric evidence that the Maya consumed amphibians (Hamblin 1984; Teeter 2001). Several authors have suggested that these animals might have been associated with shamanistic rituals and that a poison secreted by the marine toad (Rhinella marina) may have been used as a hallucinogenic drug (Hamblin 1984:53–57; Emery 2010). Because of their rarity at archaeological sites and their tendency to dig burrows, many of the amphibians recovered from Maya sites are believed to be intrusive (Wing and Scudder 1991; Carr and Fradkin

2008; Emery 2010).

Many reptile taxa had both an economic and ceremonial role for the Classic and

Postclassic Maya. Crocodiles (Crocodylus sp.) were seen as lords of the underworld

(Tozzer and Allen 1910; Emery 2010). They were offered in sacrifices and are frequently depicted in the Maya codices (Tozzer 1941; Pohl 1983:163; Thurston 2011). Snakes, associated with water and fertility, held important ceremonial role and were often depicted in scenes of blood sacrifice in Classic Maya art (Pohl 1983; Emery 2010).

Iguanas, either of the green (Iguana iguana) or black (Ctenosaura similis) variety, were prized for food in Postclassic and historic times (Tozzer 1941:191; Stuart 1964; Hamblin

1984:68–69; Emery 2010). They are often represented in the codices as sacrificial offerings (Tozzer and Allen 1910; Emery 2010:318). It is difficult to form conclusions about the use of these reptiles during the Middle Preclassic because their remains are seldom recovered in archaeological deposits from this time period.

Ethnohistoric accounts indicate that both freshwater and marine turtles were valued for their meat and eggs, as well as their carapaces, which could be used as rattles, drums, shields, and containers (Tozzer and Allen 1910; Tozzer 1941:114, 192; Stuart

1964; Emery 2010). Turtles represent one of the most abundant taxa in Middle Preclassic assemblages and, therefore, it has been suggested that the Preclassic Maya subsisted heavily on turtles (Shaw 1991; Stanchly 1995; Powis et al. 2002; Fradkin and Carr 2003).

Common species include the mud turtles (Kinosternon spp.), giant musk turtle

(Staurotypus triporcatus), common slider turtle (Trachemys scripta) and Central

American river turtle (Dermatemys mawii).

Fish probably constituted another source of proteins and fats for the Preclassic

Maya (Tozzer 1941:190–191; Hamblin 1984). Freshwater fish, such as the catfish

(Ictaluridae), mojarra (Cichlasoma spp.) and swamp-eel (Synbranchus marmoratus), were possibly locally available at most Maya sites (Powis et al. 1999; Powis et al. 2002;

Fradkin and Carr 2003; Carr and Fradkin 2008; Emery 2010). During the Middle

Preclassic, marine fish, including the bonefish (Albula vulpes), parrotfish (Scaridae), snapper (Lutjanidae) and jack (Carangidae), were sometimes imported to inland sites over considerable distances (>100 km) (Powis et al. 1999; Powis et al. 2002). During the

Classic period, marine fish bones and stingray spines were used in ceremonies performed by the elite, particularly for bloodletting rituals (Pohl 1983; Emery 2010).

Freshwater molluscs commonly found at archaeological sites included the gastropods apple snail (Pomacea flagellata) and jute (Pachychilus spp.), as well as the pearly mussel (Nephronaias spp.). Although they are occasionally eaten by the modern

Maya (Moholy-Nagy 1963, 1978; Healy et al. 1990), the frequency with which freshwater molluscs were consumed by the ancient Maya is debated (Covich 1983;

Stanchly 1995; Solis 2011). Their low nutritional returns suggest that they served at best as a supplementary source of food (Moholy-Nagy 1978; Powis 2004; Solis 2011). In archaeological settings, freshwater shells are often fashioned into utilitarian and ornamental artifacts (Moholy-Nagy 1963, 1978; Healy et al. 1990; Stanchly and Dale

1992; Harrigan 2004; Emery 2010). Frequently recovered in caches and burials from the

Late Preclassic onwards, they might have played an important role in ceremonies and mythology (Moholy-Nagy 1963, 1978; Harrigan 2004).

Marine molluscs were considered symbols of death and rebirth and often used for ceremonial and ornamental purposes from the Late Preclassic to Postclassic periods

(Moholy-Nagy 1963; Pohl 1983; Moholy-Nagy 1985; Hohmann 2002). During the

Classic period, the most highly valued ceremonial molluscs were the colorful spiny oyster

(Spondylus spp.), olive (Oliva spp.), conch shell (Strombus spp.) and marginella (Prunum apicinum) which were often worked into jewelry and decorative objects (Tozzer 1941;

Moholy-Nagy 1985; Harrigan 2004; Emery 2010). During the Middle Preclassic, marine shells were imported from the coast to inland sites for the manufacture of ornaments (Lee and Awe 1995; Hohmann 2002). It is not clear if the animal was consumed in the process

(Powis et al. 1999).

2.2.3 Middle Preclassic use of animals

The dominant foraging strategy during the Middle Preclassic involved hunting, fishing, and collecting animals procured in the immediate vicinity of the sites. Animal use included the exploitation of terrestrial animals, particularly white-tailed deer, brocket deer, armadillo, peccary, agouti, rabbit, and opossum (Pohl 1990; Shaw 1991; Wing and

Scudder 1991; Stanchly 1995; Moholy-Nagy 1998; Powis et al. 1999; Masson 2004a,

2004b; Carr and Fradkin 2008; Emery 2010) (Table 2.2). In northern Belize, the Middle

Preclassic Maya also seem to have relied on the exploitation of mud and musk turtles, which they likely acquired in the wetland areas surrounding the sites (Wing and Scudder

1991; Fradkin and Carr 2003; Masson 2004a, 2004b; Carr and Fradkin 2008).

Archaeological evidence suggests that dogs were likely consumed as food (Pohl 1990;

Shaw 1991; Wing and Scudder 1991; Clutton-Brock and Hammond 1994; Masson

2004a). At Cuello, the average mortality age of dogs (around one year old) coupled with a high frequency of spiral fractures on green bone and cutmarks support the theory of regular consumption of dogs by the Preclassic Maya (Clutton-Brock and Hammond

1994).

The Preclassic Maya likely practiced a modified form of “garden hunting”

(Linares 1976), which involved the procurement of animals near milpas. Most of the animals frequently identified in faunal assemblages are known to visit open clearings and cultivated fields (Pohl 1990; Masson 2004a, 2004b; Carr and Fradkin 2008; Emery 2010).

The presence of the brocket deer, tapir, and paca—animals that thrive in undisturbed dense forests—also indicates that hunting trips were occasionally taken to forested areas

(Pohl 1990; Shaw 1999; Masson 2004a; Carr and Fradkin 2008).

Table 2.2 List of identified taxa recovered from Middle Preclassic deposits at southern lowland Maya sites (NISP counts).

Northern Belize Central Belize Petén Scientific Name Common Name Altar de Cuello Colha K’axob Cahal Pech Bayak Seibal Sacrificios MAMMALS Didelphis spp. Opossum 69 7 26 2 Dasypus novemcinctus Armadillo 1007 11 298 4 Canidae Dog, fox, coyote 10 12 11 Canis lupus familiaris Domestic dog 381 34 17 4 9 Urocyon cinereoargenteus Gray fox 15 Procyon lotor Raccoon 16 2 Mustela frenata Long-tailed weasel 4 1 Galictis sp. Grison 2 Felidae Felines 4 1 Tapirus bairdii Tapir 1 Artiodactyla 6 1 1 Cervidae Cervids 261 27 4 5 1 Odocoileus virginianus White-tailed deer 698 68 17 31 10 13 28 Mazama americana Red brocket deer 78 2 1 2 3 Tayassuidae Peccaries 42 1 5 6 Pecari tajacu Collared peccary 5 3 Tayassu pecari White-lipped peccary 1 1 Rodentia 44 59 15 2 Dasyproctidae/Cuniculidae Agouti and paca 1 1 2 2 1 Cuniculus paca Paca 23 7 Dasyprocta punctata Agouti 32 3 2 1 Geomyidae Pocket gophers 19 Orthogeomys hispidus Hispid pocket gopher 2 5 Sylvilagus spp. Rabbits 152 2 13 BIRDS Galliformes 1 Meleagris ocellata Ocellated turkey 6 2

Northern Belize Central Belize Petén Scientific Name Common Name Altar de Cuello Colha K’axob Cahal Pech Bayak Seibal Sacrificios Other birds 10 Unidentified birds 17 6 54 39 2 AMPHIBIAN Ranidae Frogs 4 1 Rhinophrynus dorsalis Mexican burrowing toad 29 Rhinella marina Marine toad 32 REPTILES Testudines Turtles Dermatemys mawii Central American river turtle 3 12 1 13 10 Emydidae Pond turtles 88 160 Trachemys scripta Terrapin/bokatora 70 63 1 16 4 Rhinoclemmys areolata Furrowed wood turtle 103 5 2 Kinosternidae Mud and musk turtles 31 2 Claudius angustatus Narrow-bridged musk turtle 15 Kinosternon spp. Mud turtles 46 182 2 Staurotypus triporcatus Mexican giant musk turtle 68 4 1 8 2 Chelydra serpentina Common snapping turtle 708 6 16 3 Serpentes Snakes Colubridae Colubrids 8 Viperidae Vipers 6 3 Crotalinae Rattle snakes 4 Iguanidae Iguanas 6 1 2 Crocodylidae Crocodiles 3 41 2 1 1 FISH Marine fish Ariidae Sea catfish 2 Albula vulpes Bonefish 4 Carangidae Jack 5 Gerreidae Marine mojarra 4 Gobiomorus dormitor Bigmouth sleeper 3 Lutjanidae Snappers 3 1

Northern Belize Central Belize Petén Scientific Name Common Name Altar de Cuello Colha K’axob Cahal Pech Bayak Seibal Sacrificios Scaridae Parrotfish 6 Scarus spp. Parrotfish 1 Sparisoma spp. Parrotfish 18 Serranidae Grouper 1 1 Freshwater fish Ictaluridae Catfish 13 8 3 6 Cichlidae Cichlids 37 2 Cichlasoma spp. Cichlid (mojarras) 19 Synbranchus marmoratus Swamp eel 30 Unidentified fish 532 633 643 2076 80 MOLLUSCA Freshwater shellfish Pachychilus spp. Jute 3 – 3327 – Pomacea flagellata Apple snail 9 – 418 813 – 399 58 Psoronaias spp. River clam – – 3 14 Nephronaias ortmanni Pearly mussel 4 – 315 4085 – 3 Marine shellfish Strombidae Strombus – 1054 – Strombus spp. Queen conch 1 – 8 68 – Prunum spp. Marginella – 4 – Oliva spp. Olive – 2 – Dentalium spp. Tusk shell – 22 –

Source: Cuello (Miksicek 1991; Wing and Scudder 1991; Clutton-Brock and Hammond 1994; Carr and Fradkin 2008), Colha (Shaw 1999), K’axob (Harrigan 2004; Masson 2004a), Cahal Pech (Stanchly and Dale 1992; Stanchly 1995; Powis et al. 1999), Altar de Sacrificios (Pohl 1990), Bayak (Emery 2010), Seibal (Feldman 1978; Pohl 1990). – No data available.

Freshwater molluscs are not particularly abundant at Middle Preclassic sites, with the exception of Cahal Pech. These molluscs might have served as a supplementary source of food (Miksicek 1991; Stanchly 1995; Powis et al. 1999; Harrigan 2004), although their importance to the Maya diet is difficult to assess because they were not considered in the faunal analyses of Colha, Altar de Sacrificios, and Seibal. These animals could also have been used in the production of shell ornaments (Wing and

Scudder 1991). Similarly, the interpretation of fish remains is limited by the difficulty of identifying tropical fish taxa (Stanchly 1995; Powis et al. 1999). Among these, it should be noted that freshwater fish (e.g., cichlids, catfish) could have been exploited within a radius of 10 km at all sites (Shaw 1991; Wing and Scudder 1991; Masson 2004a; Emery

2010). The near absence of fish bones at Seibal and Altar is unexpected, because both sites sit on the banks of the Pasión River. This pattern may result from the recovery strategy adopted during the excavations, as sediments were not sieved at the site (Pohl

1990).

The minor representation of marine fish (e.g., bonefish, parrotfish, jack, and snapper) suggests that animal products were sometimes procured from considerable distance, but they do not seem to have been dietary staples at the sites (Shaw 1991; Wing and Scudder 1991; Stanchly 1995; Fradkin and Carr 2003; Masson 2004a; Carr and

Fradkin 2008). The inhabitants of Cuello, Colha, and K’axob may have acquired fish from Chetumal Bay (35 km to the north) or the Caribbean Sea (50 km to the east) (Wing and Scudder 1991; Fradkin and Carr 2003; Masson 2004a; Carr and Fradkin 2008). The presence of fish at the inland site of Cahal Pech indicates transport over a distance of at least 100 km (Stanchly 1995; Powis et al. 1999; Powis et al. 2002). Fish may have been

27 smoked or salted for inland transport, or kept alive in containers filled with water

(Stanchly 1995; Powis et al. 1999; Powis et al. 2002).

Likewise, marine molluscs (e.g., olive, marginella, thorny oyster, and conch shells) represent a very small component of the assemblages, with the exception of Cahal

Pech. It has been suggested that marine shells were acquired for the manufacture of artifacts (Wing and Scudder 1991; Lee and Awe 1995; Stanchly 1995; Powis et al. 1999), which might have involved consumption of the associated meat (van der Merwe et al.

2000). The presence of marine species at Middle Preclassic sites is frequently attributed to the direct exploitation of the coastal environment or the existence of exchange networks (Shaw 1991; Stanchly 1995; Powis et al. 1999; Powis et al. 2002). Animals from remote habitats are nearly absent from the sites in the Petén region (Altar de

Sacrificios, Bayal, Seibal, and Tikal), with the exception of a stingray spine recovered from a burial at Tikal (Moholy-Nagy 1998).

2.2.4 Stable isotope analysis

Animal and plant remains recovered from archaeological sites are useful for determining the types of resources available to the ancient Maya. Stable isotope analysis can add to these results by determining which foods were consumed and in what proportions. More precisely, stable isotope analysis of human bones can help detecting the consumption of maize (a C4 plant) in comparison to C3 plants (e.g., root crops, fruits, nuts, legumes, and vegetables), as well as determining the main sources of proteins

(terrestrial versus aquatic resources) in the diet.

Stable isotope studies from Middle Preclassic sites suggest considerable dietary heterogeneity between sites located in Belize versus the Petén region (White and

Schwarcz 1989; Tykot et al. 1996; White 1997; Wright 1997, 2006). Such variation has been attributed to differences in local ecology and population density (van der Merwe et al. 2000; Wright 2006:196). People at sites located in the Petén (i.e., Seibal and Altar de

Sacrificios) are believed to have relied heavily on maize and animals characterized by a diet rich in C4 plants. These include domestic dogs and animals that may browse in agricultural fields, such as deer and peccaries (Wright 1997, 2006). The high nitrogen values observed in the analyses may be indicative of the consumption of large quantities of terrestrial herbivores as well as freshwater fish (Wright 2006:192–196).

In comparison, the Preclassic Maya living in northern Belize apparently subsisted on a broader diet, consuming a wider range of plant foods and animal resources (van der

Merwe et al. 2000; Wright 2006). Consistently more negative collagen values at Belizean sites suggest a greater access to plants other than maize relative to the Petén region

(White and Schwarcz 1989; Henderson 1998; Coyston et al. 1999; Henderson 2003). The range of carbon and nitrogen values may indicate that a variety of terrestrial animals, such as deer, dog, and peccary, constituted the main protein sources, with the possible addition of turtles (White and Schwarcz 1989; Tykot et al. 1996; Henderson 1998). The stable isotope analyses reveal that marine and reef resources do not seem to have contributed substantially to the diet in northern Belize. This result is not unexpected considering that all three sites are located approximately 50 km inland (White and Schwarcz 1989;

Henderson 1998; van der Merwe et al. 2000; Young 2002; Hammond and Young 2003;

Henderson 2003). Small amounts of freshwater fish, however, were presumably consumed at Cuello (Young 2002:117–118; Hammond and Young 2003) and Lamanai

(White and Schwarcz 1989).

Located only seven km from the Caribbean Sea, the residents of Altun Ha seem to have relied heavily on the exploitation of the coastal environment. Stable isotope analyses suggest that marine and reef resources likely constituted the main protein sources at the site (White et al. 2001a). This interpretation is supported by the recovery of many vertebrate and molluscan marine remains (Pendergast 1979:7–12). Maize is argued to have formed the main dietary staple, with the addition of other wild and cultivated plants

(White et al. 2001a).

At Cahal Pech, in central Belize, the carbon and nitrogen isotopic values are similar to those observed at inland sites in northern Belize. This further supports the argument that maize represented a significant, but not dominant, component of the diet during the Middle Preclassic (Powis et al. 1999). Combined carbon and nitrogen values suggest that terrestrial herbivores were consumed as well as reef fish. This last result is surprising because the frequent consumption of marine fish seems incompatible with the location of the site (Powis et al. 1999).

2.3 Summary

Faunal, paleobotanical, and isotopic data recovered from sites in the southern

Maya lowlands points to a broad-based subsistence pattern during the Middle Preclassic period. The ancient Maya were relying heavily on cultivated plants, such as maize, squash, and beans, which they supplemented with an assortment of fruits, root crops, and vegetables. They consumed wild game and domesticated animals, as well as invertebrates and wild plants. The next chapter turns to a description of the research undertaken at

Pacbitun.

CHAPTER 3: SITE DESCRIPTION AND PREVIOUS RESEARCH

This chapter provides contextual information about the ancient Maya site of

Pacbitun. It begins with a description of the environmental setting and geographic location of Pacbitun. Then, the various archaeological excavations carried out at the site are described, followed by a presentation of the history of occupation of Pacbitun. The chapter concludes with a description of the architectural remains and artifactual assemblages dating from the Middle Preclassic period, and a discussion of local subsistence practices.

3.1 Geographical context

Pacbitun is a medium-sized center located within the southern Maya lowlands, in west-central Belize. It lies on the southern edge of the upper Belize River Valley, three kilometers east of the modern village of San Antonio (Figure 3.1). The site was occupied between 900 BC and AD 900 and is one of many centers that flourished during the

Classic period (AD 300–900) in the Belize River Valley (Healy et al. 2004b; Healy et al.

2007).

Pacbitun’s location on the southern rim of the Belize River Valley, at the junction of two ecozones, namely the lowland tropical rainforest and the upland pine ridge, made it unique from other sites in the area. It has been suggested that this location was chosen by the early Maya to take advantage of the locally contrasted microenvironments and their diverse resources (Graham 1987; Campbell-Trithart 1990; Healy 1990a). The lowland tropical rainforest borders Pacbitun to the west, north, and east. This ecozone is

Figure 3.1 Map of the Belize Valley (modified from Chase and Garber 2004:2). characterized by a dense tropical broadleaf forest which provides a habitat for numerous animals (e.g., agouti, coati, white-lipped peccary, curassow, and red brocket deer)

(Schlesinger 2001). Around Pacbitun, fertile soils are found in natural catchment basins and depressions created by the hilly terrain (Wright et al. 1959:190–191; Healy 1990a;

Sunahara 1995; Healy et al. 2007). Further south, the tropical rainforest gives way to the

Mountain Pine Ridge (Figure 3.2), an ecozone that is sparsely covered with highland oak and pine. The lack of cultivable lands in this region has limited permanent habitation by the ancient Maya, although they likely exploited other resources, such as granite, slate, and pinewood (Wright et al. 1959:171–175; Graham 1987; Healy 1990a).

Figure 3.2 Location of Pacbitun in the Upper Belize Valley (Hohmann and Powis 1999:2).

The Pacbitun Maya also had access to the Belize River and its two principal tributaries, the Mopan (Western Branch) and Macal (Eastern Branch) Rivers. In addition, multiple secondary and tertiary streams are found within kilometers of the site, such as

Barton Creek, Slate Creek, Tutu Creek, and Privassion Creek (Figure 3.2) (Campbell-

Trithart 1990; Hohmann and Powis 1999). This river system provided the ancient Maya with continuous access to aquatic resources, such as fish, molluscs, waterfowl, iguanas, and turtles (Hohmann 2002:61), and facilitated the transportation of goods from the

Caribbean Sea to the interior of the Maya subarea (Willey et al. 1965; Chase and Garber

2004). Settlement surveys at Pacbitun also revealed the existence of permanent water springs and a major reservoir near the site (Healy et al. 2007).

Pacbitun is located within the tierra caliente (“hot land”) zone of the tropics (West and Augelli:46). In this tropical to sub-tropical climate, temperatures are nearly constant throughout the year, ranging between 25–30°C (Wright et al. 1959; West and Augelli

1966). The climate is also characterized by distinct dry and wet seasons. In the San

Antonio sub-region where the site of Pacbitun is located, the dry season lasts from

January to April, whereas the wet season occurs between May and December. This sub- region receives about 125–175 cm of rainfall per year (Wright et al. 1959). Palaeoclimate records from the lakes of Punta Laguna (Curtis et al. 1996) and Chichancanab (Hoddell et al. 1995), in northern Yucatan Peninsula, suggest that the climate during the Middle

Preclassic was slightly wetter than today.

The central precinct of Pacbitun lies 240 meters above sea level and sits atop a limestone plateau that is oriented east-west. This elevated position in the landscape provided a peripheral view of the hilly terrain surrounding the site, which can be divided into three zones: the epicenter, the core zone, and the periphery (Figure 3.3). Research suggests that the epicenter (0.5 km2) formed the religious and political heart of Pacbitun during the 2000 years of occupation of the site (Healy et al. 2007:17–18). The current configuration of this zone consists of 40 masonry structures, including temple-pyramids, palace-like range structures, one ball court, two causeways, and 20 whole and partial stelae and altars (Figure 3.4). It also contains three major (A, B, and C) and two secondary (D and E) plazas, the latter located to the north of the main site axis, and a number of smaller courtyard groups. During the Classic period, the use of the epicenter was likely restricted to the elite class, with few exceptions (Bill 1987; Healy 1990a;

Healy et al. 2007).

Epicenter

Core Zone

Periphery Zone

Figure 3.3 Map of the Pacbitun settlement survey. The concentration of monumental architecture in the center of the map forms the epicenter. The dashed square around the epicenter coincides with the 1 km2 core zone. The four dashed rectangles delineate the periphery zone. All architectural remains are drawn in black. Shaded circular areas are possible reservoirs (modified from Healy et al. 2007:20).

The core zone includes the epicenter as well as small, medium, and large mounds constructed within an area of 1 km2 from Plaza A (Figure 3.5). The earthen mounds are the remains of domestic structures that are generally considered residences for lesser-elite or non-elite individuals living outside the epicenter (Campbell-Trithart 1990:319–322;

Healy et al. 2007). The periphery zone surrounds the core zone and covers an additional 8 km2 (Figure 3.3). This zone was probably the sustaining agricultural area of Pacbitun. It is

Figure 3.4 Plan of the epicenter of Pacbitun. Structures are shown as first found (unexcavated) and rendered in the standard isometric convention. Stelae and altars are drawn as conventional rectangles and circles, respectively (modified from Healy et al. 2007:19). characterized by a scatter of small housemounds, typically located on hilltops or elevated ridges, perhaps in order to maximize the use of fertile lands for agriculture (Richie 1990;

Sunahara 1995:132–133; Healy et al. 2007). Most structures were probably residential in nature, but some may have been used for small-scale craft production, storage, or as kitchens (Richie 1990:199; Sunahara 1995:103; Healy et al. 2004b).

3.2 Archaeological investigations

The first reference to the site of Pacbitun is attributed to D. H. Snow (1969), an ethnologist conducting research in San Antonio, who mentioned the presence of

Figure 3.5 Core zone of Pacbitun, indicated by dashed lines. Core zone mounds are rendered as black-shaded forms. Possible reservoirs are represented by gray-shaded circular areas (Healy et al. 2007:27). prehistoric mounds and a large pyramidal structure—presumably Structure 1—near the village. In 1971, the site came to the attention of the Department of Archaeology of

Belize (then British Honduras) when Structure 41 was quarried for the construction of a modern roadway. Peter Schmidt, then Archaeology Commissioner, conducted a preliminary survey and officially designated the site as “Pacbitun,” which translates to

“stones set in earth.” The name might refer to the standing stelae in Plaza A (Healy

1990a).

A surface survey was conducted in 1980 by Paul F. Healy. This work showed the presence of numerous well-preserved architectural structures and a widely terraced periphery. Systematic archaeological excavations were first conducted at Pacbitun in

1984, 1986 and 1987 by the Trent University-Pacbitun Archaeological Project, directed by Healy. The aim of the project was to provide a comprehensive perspective on the ancient occupation of Pacbitun by documenting the diachronic development of the site, both in the site core and its periphery (Healy 1990a). Excavations in the epicenter exposed the different architectural phases of constructions, including those relevant to

Structures 1, 2, 4, 5, 14, 15, and 23 (Bill 1987; Healy 1988, 1990a, 1992; Healy et al.

2004a; Healy et al. 2004b). The survey of the core zone focused on an area of 1 km2 around the epicenter (Campbell-Trithart 1990), while four transect zones extending 1 km from the epicenter mapped the housemounds located in the periphery zone (Richie 1990;

Sunahara 1995).

Excavations were also conducted from 1995 to 1997 by the Trent University-

Preclassic Maya Project in Plazas B, C, and D (Arendt et al. 1996; Hohmann and Powis

1996, 1999; Hohmann et al. 1999). This project aimed to document the Preclassic occupations at the site, with a focus on the organization, social structure, subsistence, and economy of Middle Preclassic domestic households (Healy and Awe 1995a, 1995b, 1996;

Healy 1999c). The research also examined the evidence for specialized shell production and its role in the development of socioeconomic complexity in the Belize River Valley

(Hohmann and Powis 1996, 1999; Hohmann et al. 1999; Hohmann 2002).

Since 2008, new excavations in Plazas A and B were performed by the Pacbitun

Regional Archaeological Project (PRAP), under the direction of Terry G. Powis (Powis

2009, 2010, 2011). This project focuses on documenting the earliest occupations at

Pacbitun. In addition, PRAP also investigates caves, causeways, and minor centres in the southern periphery of Pacbitun (Powis 2010; Spenard 2011; Valdez et al. 2011; Weber

2011; Weber and Powis 2011; Powis 2012b). This new research program seeks to define the extent of settlement around the site and to document the interactions between

Pacbitun and its periphery (Powis 2010:23).

During the 1995–1997 and 2008–2011 field seasons, excavations were conducted by trowel and shovel, following the cultural stratigraphy. Artifacts were recovered from both primary (floor and perimeter deposits) and secondary (construction fill, structural fill, and secondary middens) contexts. Particular attention was given to the excavation of

Plaza B sub-structures in 2008–2010. Floor deposits were excavated according to 5 cm intervals in order to control for the recovery of artifacts embedded in the floor surface.

Any artifacts found below the initial 5 cm depth were considered as secondary fill rather than primary floor deposits. During the excavations, visible artifacts were collected by hand and all deposits were dry-screened in the field using a 1/4 inch (6 mm) wire mesh screen. Additionally, in 2008 and 2009, all the floor deposits from Sub-Structures B-1 and

B-2 in Plaza B were wet-screened with the use of 1/16 inch (1.2 mm) mesh screen (Powis

2009:18–20).

3.3 Site chronology

Excavations at Pacbitun have revealed a long sequence of occupations extending from the early Middle Preclassic (ca. 900 BC) to the Terminal Classic (ca. AD 900)

39 periods (Healy 1990a). The cultural history was first established by cross-dating ceramics with cultural sequences already established for other sites in the Belize Valley (Figure

3.6), specifically Barton Ramie (Gifford 1976) and Cahal Pech (Awe 1992). The chronology was then tested with 22 radiocarbon dates (Healy 1990a, 1999b; Healy et al.

2004b). Additional radiocarbon dates obtained by PRAP also helped to refine the chronology for the Middle Preclassic occupations (Powis 2009).

Figure 3.6 Chronology and ceramic complexes of Pacbitun in comparison to those of Barton Ramie, Xunantunich, and Cahal Pech (Healy et al. 2007:21).

To this date, there has been no secured evidence that Pacbitun was permanently or densely occupied before the Middle Preclassic (900–300 BC). This time period is locally

40 referred to as the Mai phase and is subdivided into early and late periods at ca. 650 BC.

Pacbitun was then a small farming village, relying on swidden agriculture (Wiesen and

Lentz 1999) and involved in the manufacture of shell ornaments (Hohmann 2002; Powis

2009, 2010). The village was likely confined to the epicenter given that surveys of housemounds in the core and periphery zones failed to find evidence of clear Middle

Preclassic occupation (Sunahara 1995:108; Healy et al. 2007). Because the Middle

Preclassic is the focus of the present study, this period is discussed in greater details in sections 3.4–3.6.

The Late Preclassic Puc (300–100 BC) and Terminal Classic Ku (100 BC – AD

300) phases witnessed the construction of the first monumental architecture in the site core. The major buildings in Plaza A formed an E-Group complex, which consisted of a standard trio of temple-pyramids (Structures 1, 4 and 5) opposing a lone temple-pyramid

(Structure 2) (Figure 3.4). Plazas A and E were plastered during this time period and a ceremonial ballcourt (Structures 14 and 15) was also constructed (Healy 1990a, 1992;

Healy et al. 2004b). The presence of E-Groups and ballcourts at Pacbitun likely reflects an early involvement in ceremonial activities at the site (Healy 1990a, 1992). Importantly, it is during this time period that the core zone and the periphery were first settled by small groups (Campbell-Trithart 1990:313; Richie 1990:194–195; Sunahara 1995:100; Healy et al. 2007).

During the Early Classic Tzul phase (AD 300–550), many major public structures were enlarged (e.g., Structures 1, 2, 4, 5, 14, and 15), while others, such as elite residences (Str. 23 and 38), were constructed (Healy 1990a; Cheong 2013). Elite individuals were interred with substantial quantities of exotic goods (Bill 1987; Healy

1990a). New monuments were erected, including a carved stela (Stela 6) portraying the

41 accession to the throne of a Pacbitun lord around AD 485. This stela is one of the earliest dated monuments in the southern Maya lowlands (Healy 1990b; Helmke et al. 2006). All these activities attest to the increasing wealth of the site and suggest that Pacbitun was already playing a dominant role in the eastern region of the southern lowlands during this period (Healy et al. 2004b).

The developmental peak of Pacbitun occurred during the Late Classic Coc phase

(AD 550–700). Almost every building in the epicenter underwent massive architectural renewal at this time (P.F. Healy, 2013, personal communication). Large stone monuments were carved and erected. Fine local or imported objects were frequently deposited as offerings in elite burials (Healy 1988, 1990a; Healy et al. 2004a). The population is believed to have peaked during the Late to Terminal Classic period, with an estimate of

5000–7000 persons living within an area of 9 km2. This coincided with an unprecedented period of agricultural intensification, resulting in the widespread use of hillslope terracing

(Campbell-Trithart 1990:317–318; Richie 1990:194–197; Healy 1990a; White et al. 1993;

Sunahara 1995:114–116; Healy et al. 2007). The periphery was extensively occupied during this time period (Richie 1990:198–199; Sunahara 1995:100).

The site continued to grow and expand during the Terminal Classic Tzib phase

(AD 700–900), although on a smaller scale, before it started to decline. Pacbitun was abandoned around AD 900. There is little evidence of activity during the subsequent

Postclassic period (Richie 1990:194; Healy 1990a).

3.4 Middle Preclassic investigations

Given that the faunal remains discussed in this study were recovered exclusively from Mai phase deposits, more information is provided about the Middle Preclassic

42 architectural and artifactual findings of the 1995–1997 and 2008–2011 excavations at

Plaza A, B, C, and D (Figure 3.4).

The earliest architectural remains of Plaza B date to the early Middle Preclassic

(900–650 BC) and consist of two partially exposed platforms (Sub-Structures B-1 and B-

4) and a wall (Sub-Stone Alignment B-12) lying about 10–15 centimeters above the bedrock (Figure 3.7). The structures were modest, consisting of low platforms with retaining walls formed of two courses of roughly-shaped limestone blocks. The presence of postholes in the decomposed bedrock suggests that the platforms supported perishable superstructures (Hohmann and Powis 1996, 1999; Hohmann et al. 1999). These platforms are the earliest constructed at Plaza B and, perhaps, at Pacbitun. Two radiocarbon samples collected from Sub-Str. B-1 provided an age of 815–530 BC and 760–410 BC3. Those dates are consistent with the ceramic typology (Healy 1999b; Powis, personal communication, 2012).

Some changes occurred during the late Middle Preclassic (650–300 BC). Sub-Strs.

B-1 and B-4 were abandoned around 650 BC and partially covered over, providing a stable foundation for the construction of eleven new platforms (Sub-Strs. B-2, 3, 5–11,

13, and 14). Radiocarbon samples retrieved from Sub-Strs. B-2 and B-3 provided date ranges of 770–375 BC and 905–400 BC, respectively (Healy 1999b). The newer platforms were better constructed and larger than previous phases of construction, with walls made of three courses of cut limestone blocks (Hohmann and Powis 1996). The close proximity of the platforms and common extramural areas suggest that the structures were organized as a patio group with several platforms situated around an open plaza.

3 All radiocarbon dates provided in this study are calibrated and presented with two sigma deviations.

Figure 3.7 Plan of the Middle Preclassic sub-structures in Plaza B. The early Middle Preclassic structures are shaded in black (Hohmann et al. 1999:22).

This is a common residential pattern observed at many lowland Maya sites (McKillop

2004a:150; Powis 2010:11). While each of the buildings may have served a distinct function, such as kitchens, storage areas, or residences, the presence of substantial quantities of domestic refuse both within and around the structures suggest that they were principally domestic households (Hohmann and Powis 1999; Powis 2010).

Sometime during the late Middle Preclassic, the late Mai sub-structures were abandoned and covered with a dense, midden-like deposit. The midden is characterized by a dark organic soil with significant quantities of artifacts and ecofacts. Because it was

44 not found in association with architectural features, it has been inferred that the midden was made from materials derived from other areas of the site and used as construction fill to level and possibly enlarge the plaza (Hohmann and Powis 1999). A radiocarbon sample provided a date range of 780–395 BC for this construction (Healy 1999b), whereas the ceramic typology indicates that it was likely deposited closer to the end of the late Mai phase, ca. 400–300 BC (Hohmann and Powis 1996). Despite the discrepancy in the age estimates, both sets of dates fall within the late Middle Preclassic period.

Excavations in Plaza A have exposed an alignment dating from the transitional period (300–100 BC) between the late Middle Preclassic and early Late Preclassic.

Located in front of Str. 4, this wall made of cut limestone possibly forms the edge of a platform, but further excavations are needed to determine its size and function. The modes of construction of this sub-structure are very similar to the sub-structures found in

Plaza B (T. G. Powis, personal communication, 2012). In the center of the plaza, a retaining wall (Sub-Str. A-1) was also uncovered (Figure 3.8). Ceramics found outside of the structure, below a plaza floor, provided a late Middle Preclassic date (400–200 BC) for this construction. Below this floor, three additional sealed plaza floors (Levels 6–8) were uncovered. An early Middle Preclassic occupation was confirmed by a radiocarbon sample from Level 8 (which provided a date range of 700–400 BC) and the ceramic typology. Four plaza floors with a comparable chronology were also observed inside Sub-

Str. A-1 (Powis 2011; T. G. Powis, personal communication, 2012). The chronology for

Plaza A shows the same temporal sequence identified in Plaza B (T. G. Powis, personal communication, 2012) and on the adjacent Eastern Court (Cheong 2013).

Excavations in Plaza C have revealed the presence of two early Mai (Sub-Str. C-2 and C-3) and two late Mai structures (Sub-Str. C-1 and C-4) (Arendt et al. 1996;

Hohmann et al. 1999; Hohmann 2002:191). Two additional Middle Preclassic stone alignments were uncovered on the western slope of Plaza C (Hohmann et al. 1999). These structures are all similar to those uncovered in Plaza B, although no alleyway or perimeter deposit were encountered (Hohmann 2002:191). A cist burial (BU-C1) dating to the

Middle Preclassic was also identified. The interment included a 30–40 years old individual lying in a prone position, with the head oriented to the west. The cist contained one mano fragment, two shell disk beads, one obsidian blade fragment, and three simple vessels (Arendt et al. 1996; Healy et al. 2004b).

Figure 3.8 Plan of the Middle Preclassic Sub-Structure A-1 in Plaza A. Image courtesy of PRAP.

A small test unit in Plaza D exposed a total of five marl floors and four stone alignments (Sub-Stone Alignments D-1, 2, 3, and 4). The earliest stone alignment dates to the transition from the early to late Middle Preclassic, ca. 650–600 BC. The three others date to the late Middle Preclassic. Artifacts were sparse (Hohmann et al. 1999).

3.5 Artifactual assemblages from the Middle Preclassic period

The artifacts recovered at Pacbitun are numerous and diverse. Local materials include granite, slate, and chert (Table 3.1). These could be quarried from sources in the

Belize River Valley as well as the Mountain Pine Ridge (Figure 3.2) (Graham 1987;

Healy et al. 1995; Hohmann and Powis 1996). Three types of Preclassic artifacts were acquired through long-distance trade: obsidian, jade, and marine shell (Table 3.1).

Physiochemical analysis of 18 Middle Preclassic obsidian specimens demonstrated that the volcanic glass was imported from three highland Guatemalan sources (Healy 1990a;

Awe et al. 1996; Hohmann and Powis 1996:118–119), while the greenstone and jade possibly came from the Motagua River Valley of Guatemala (Healy 1990a; Hohmann and

Powis 1996).

Marine shells were imported whole from the Caribbean coast and used in the manufacture of shell ornaments. A small proportion of the artifacts were also made from local freshwater species, namely the pearly mussel and the locally available jute

(Hohmann 2002:115–117; Powis 2009, 2010). The majority of the shells were transformed into small disks and irregular beads (Hohmann 2002:125). Evidence suggests that the early Maya of Pacbitun were involved in the specialized production of shell ornaments at the household level. The majority of ornaments were likely destined for

47 export to other Maya sites, likely into the Petén region of Guatemala (Hohmann

2002:192–194; Powis 2009).

Table 3.1 Possible source location for local and exotic raw materials found at Pacbitun.

Artifact types Local/Exotic Possible sources Distance Granite Local Stann Creek 5–10 km Macal River Mountain Pine Ridge

Slate Local Slate Creek 5–10 km Little Vaquero Creek Barton Creek

Chert/Chalcedony Local Belize River Valley 10–20 km

Marine shell Exotic Caribbean coast ~100 km

Jade/Greenstone Exotic Motagua River Valley ~150 km

Obsidian Exotic El Chayal 400–500 km San Martin Jilotepeque Ixtepeque

Temporal changes in the quantity and quality of artifacts recovered from the sub- structures suggest subtle social and economic differences during the Middle Preclassic

(Arendt et al. 1996; Hohmann and Powis 1996, 1999; Hohmann et al. 1999; Powis 2009,

2010). The artifacts indicate that the Pacbitun inhabitants were essentially egalitarian during the early Middle Preclassic. During the late Middle Preclassic, the social system may have become gradually hierarchical, as indicated by the higher frequency and greater diversity of long-distance exchange goods and the presence of substantial quantities of shell ornaments. There are also signs of a greater degree of standardization in the production of artifacts, such as disk shell beads (Hohmann 2002).

3.6 Subsistence practices

As discussed in Chapter 2, it is believed that the Preclassic Maya subsisted primarily on the cultivation of three cultigens: maize, beans, and squash (Turner and

Miksicek 1984; Lentz 1999; Lentz et al. 2005). Unfortunately, evidence for agriculture during the Middle Preclassic is scarce at Pacbitun, perhaps as a result of poor organic preservation. Few preserved kernel cupules of maize were identified in two late Middle

Preclassic samples (Wiesen and Lentz 1999), but no paleobotanical remains of beans or squash were recovered. The recovery of multiple fragments of grinding tools, such as mano and metate, may constitute indirect evidence for the consumption of wild and domesticated plants, including maize (Hohmann and Powis 1996; Duffy 2011). Remains of the coyol palm, “turtlebone” (Pithecolobium sp.), and ramón have also been identified

(Wiesen and Lentz 1999). Wiesen and Lentz (1999) suggest that the residents of Pacbitun possibly exploited these plants for various usages, such as food, medicine, artifact manufacturing, construction materials, and fuel.

Faunal remains recovered from Preclassic contexts during the 1995 and 1996 excavations in Plaza B and C were previously analyzed by Norbert Stanchly (1999). The preliminary report provided NISP counts (Table 3.2). MNI values were not calculated.

According to Stanchly (1999), a total of 2,327 specimens were recovered, with 1,575 invertebrate remains (67.7% of the assemblage) and 752 vertebrate remains (32.3%).

Freshwater molluscs include jute snails, apple snails, and pearly mussels. More than 100,000 jute remains were recovered between 1995 and 1997 from Plaza B investigations. From this, 225 jute were selected for analysis. All specimens presented a broken apex or punctured spire. Both methods are believed to facilitate the detachment of the snail from the shell after it is cooked (Healy et al. 1990; Stanchly 1999). Based on the

Table 3.2 Vertebrate and invertebrate remains identified by Stanchly (modified from Stanchly 1999:49–50).

VERTEBRATES INVERTEBRATES Taxon n Taxon n Osteichthyes Gastropoda Ictaluridae 1 Strombus pugilis 2 Unidentified fish 1 Strombus sp. 565 Aves Strombus sp.? 4 Bird? 12 Pachychilus indiorum 158 Unidentified bird 1 Pachychilus glaphyrus 64 Reptilia Pachychilus sp. 3 Chelonia? 6 Pachychilus sp.? 1 Chelonia 3 Pomacea flagellata 24 Kinosternon spp. 14 Pomacea sp.? 4 Staurotypus triporcatus? 1 Prunum apicinum 12 Unidentified reptile 2 Orthalicus sp.? 1 Mammalia Olividae 1 Dasypus novemcinctus 12 Neritida 1 Dasypus novemcinctus? 1 Pelecypoda Sylvilagus sp. 1 Nephronaias ortmanni 725 Sylvilagus sp.? 1 Donax sp. 1 Tapirus bairdii 1 Chamidae 1 Carnivora 1 Arcinella sp. 1 Canis lupus familiaris 1 Scaphopoda Canis lupus familiaris? 2 Dentalium sp. 1 Cervidae 66 Mollusca Cervidae? 29 Unidentified shell 5 Odocoileus virginianus 53 Mazama americana 3 Tayassuidae 4 Rodentia 1 Rodentia? 1 Cuniculus paca 5 Cuniculus paca? 1 Homo sapiens 4 Unidentified mammal 434 Unidentified specimens 90 Total 752 Total 1575

extremely large numbers of jute recovered at Pacbitun in fill and midden contexts, Healy and colleagues (1990), as well as Stanchly (1999), suggested that the freshwater snail possibly played a significant role in the subsistence of the Preclassic Maya. However, more recent analyses by Solis (2011) indicate that natural processes and rodent activity

50 may alter the shells in a fashion identical to the patterns created by cultural activities.

Therefore, broken spires may not necessarily be indicative of human consumption of jute, as frequently argued in the Maya literature. Solis (2011) suggested instead that the presence of jute in Terminal Preclassic deposits at the site of Minanha is more likely due to the incidental inclusion of jute in river clays used as construction fill. A re-examination of the archaeological contexts and artifacts associated with the Pachychilus remains could help to determine the use of these snails at Pacbitun.

Similarly, the importance of apple snails and pearly mussels to the diet is poorly understood (Stanchly 1999). Powis (2004) has suggested that freshwater mussels possibly served as a dietary supplement or famine food when other protein sources were unavailable. Marine molluscs, which were used for the manufacture of shell ornaments, may also have been imported from the coast as a source of food, the meat being salted or smoked (McKillop 2004b). However, data is currently lacking to test these hypotheses.

Of the 752 vertebrate remains from Plaza B and C, 224 were identified below class level by Stanchly (1999). Mammalian species include white-tailed deer, red brocket deer, paca, rabbit, armadillo, domestic dog, peccary, and tapir. Turtle remains are represented by shell fragments of small mud turtles and possibly the giant Mexican musk turtle. Fish remains include one catfish element. Bird remains are also present, but are too fragmented to be identified below class level. The size of the fragments suggests the presence of a medium to large bird. All of these vertebrate species were probably consumed as subsistence items. Deer specimens dominate the assemblage. The unidentified mammalian fragments are consistent with this picture (Stanchly 1999).

Stanchly suggests that the Preclassic Maya of Pacbitun hunted both small and large terrestrial game and gathered turtles and shellfish. Most of the animals could have been

51 obtained from local ecozones. The wide spectrum of species exploited, despite the small sample of the assemblage, is typical of lowland Maya faunas (Stanchly 1999).

3.7 The Middle Preclassic at Pacbitun: A summary

During the Middle Preclassic period, which is the focus of this research, Pacbitun was a small farming village consisting of a dispersed scatter of households, as evidenced by the presence of sub-plaza structures in the epicenter. Indeed, the various programs of excavations revealed that the Middle Preclassic settlement extended to portions of at least four plazas (A, B, C, and D), covering an area of 300 m (east-west) by 125 m (north- south). There seems to be no evidence of Middle Preclassic settlement outside the epicenter based on the extant data. Healy and colleagues (2007) have estimated a population of 16–49 persons, but this value is likely underestimated because it only takes into account the structures uncovered in Plaza B.

The various classes of artifacts recovered within and around the structures (e.g., ceramics, jade and shell ornaments, lithics tools) all indicate that the Middle Preclassic residents of Pacbitun were involved in typical household activities, such as food preparation and tool manufacture. The presence of exotic goods, such as obsidian, jade, and marine shells, confirms that Pacbitun was participating in inter-regional exchange systems at an early date. The preliminary paleobotanical and archaeozoological data suggest that the Pacbitun Maya practiced swidden agriculture, which they may have supplemented with the hunting and gathering of an array of locally available animals and plants.

This chapter has presented the geographical context and cultural history of

Pacbitun as well as detailed the archaeological investigations undertaken at the site. This

52 background information will be necessary to an interpretation of the Pacbitun faunal material (Chapter 7). The following chapter turns to the theoretical framework used to analyze this material.

CHAPTER 4: FORAGING THEORY

The theoretical approach guiding this research is foraging theory. This chapter defines and examines three types of foraging models used in archaeozoology: the prey choice model, patch choice model, and central place foraging model. After presenting the assumptions and predictions of each model, this chapter considers how the central place foraging prey choice model is most appropriate for interpreting faunal assemblages at

Pacbitun. It also discusses how foraging theory can be applied to the archaeological record. The second part of the chapter offers a description of the ecology and behavior of the species identified at Pacbitun.

4.1 Theoretical approach

Foraging theory is a family of models derived from human behavioral ecology.

Central to this theoretical framework is the premise that foraging behaviors are under selection and contribute to enhance an individual’s chances of survival and reproductive success (Smith 1983; Bird and O'Connell 2006). Specifically, in anthropology, it is assumed that foragers were concerned with maximizing their net rate of energy acquisition under a certain set of environmental conditions. This approach is commonly applied to the study of subsistence strategies, transport decisions, and technological changes (Bird and O'Connell 2006). The following section discusses three fundamental sets of foraging models used in archaeozoology to examine the decisions underlying the acquisition of animal resources.

4.1.1 Prey choice model

The prey choice model, also known as the diet breadth model, aims to predict which resources a forager should include in its diet or ignore. When foragers encounter prey, they have to decide whether they should exploit it or continue searching for a more profitable one (Smith 1983; Stephens and Krebs 1986). The prey choice model assumes a

“fine-grained” environment; that is, an environment within which prey are randomly distributed (MacArthur and Pianka 1966; Smith 1983). The model further postulates that a forager searches for all types of prey simultaneously and encounters them sequentially in the environment (MacArthur and Pianka 1966; Stephens and Krebs 1986; Bird and

O'Connell 2006; Lupo 2007). Foraging time is divided into two mutually exclusive activities: searching and handling. Searching is defined as the time devoted to looking for resources. Handling consists of the time spent pursuing, capturing, processing, and consuming a prey item that has been encountered (MacArthur and Pianka 1966; Smith

1983; Stephens and Krebs 1986; Bird and O'Connell 2006; Lupo 2007). The model postulates that foragers can accurately estimate the likely encounter and post-encounter return rate for all potential prey types because they possess prior information about the abundance of resources in a habitat and the costs and benefits associated with their acquisition (Stephens and Krebs 1986).

The prey choice model requires resources to be ranked as a function of their

“profitability”, which is measured as the net post-encounter return rate (Smith 1983;

Stephens and Krebs 1986; Bird and O'Connell 2006; Lupo 2007). According to the model, prey types are added to—or deleted from—the diet as a function of their rank order so that the overall foraging return rate is maximized (Smith 1983; Stephens and

Krebs 1986; Bird and O'Connell 2006; Lupo 2007). As a result, foragers are expected to

55 always pursue the highest-ranked prey type whenever encountered because this strategy provides the highest net gain of energy per unit of handling time (Smith 1983; Bird and

O'Connell 2006; Lupo 2007). Less profitable prey types, regardless of their abundance in the environment, should be included in or excluded from the diet depending on the probability of encountering higher-ranked resources. In fact, low-return prey taxa should always be ignored if their acquisition does not result in an increase of the overall foraging return rate (Smith 1983; Stephens and Krebs 1986).

Given these predictions, variations in diet breadth (i.e., number of prey types exploited) can be used to assess foraging efficiency. In a rich environment, foragers should have a narrow diet dominated by a relatively small number of prey types.

However, if the availability of highly profitable resources declines substantially, either through predation or environmental change, foragers may be forced to handle an increasing number of less profitable prey types. This situation should progressively produce a broader diet (Smith 1983).

The prey choice model postulates that prey types are found in a fine-grained environment. However, this assumption is violated when resources are clumped together, that is, when they form “patches.” The patch choice model, which is discussed next, represents a useful alternative in this situation.

4.1.2 Patch choice models

First developed by MacArthur and Pianka (1966) as a complement to the prey choice model, the patch choice model predicts foraging behaviors in environments where resources occur in patches (Smith 1983). Patches are assumed to be distant from one another and the time spent travelling from one patch to another is considered

56 unproductive (Stephens and Krebs 1986). Two mutually exclusive activities are defined by the model: travelling and handling (MacArthur and Pianka 1966; Lupo 2007). The patch choice model presumes that patches are encountered randomly and sequentially in the environment and are ranked according to their respective profitability (MacArthur and

Pianka 1966; Stephens and Krebs 1986). Ranks are established as a function of the return rate expected from searching for and handling prey types within each patch, while taking into account the cost of travelling to them (Bird and O'Connell 2006). Overall, the model predicts that foragers add patches to their itinerary in rank order until the average foraging return per unit (including travelling) declines (Smith 1983; Lupo 2007).

Although the patch choice model is designed to determine which patches foragers should exploit, it does not consider time allocation within these patches. The marginal value theorem proposed by Charnov (1976) acts as a natural extension to the patch choice model and focuses on explaining these issues (Smith 1983; Stephens and Krebs 1986). At the heart of the model is the assumption that foraging generally depletes a patch from its resources (Charnov 1976). As a forager spends more time in a patch, highly-ranked prey normally become rarer; the energy cost of searching for resources gradually increases, which results in a steady decline of the marginal return rate of that particular patch (Smith

1983; Winterhalder and Kennett 2006; Lupo 2007). As a consequence, the marginal value theorem predicts that a forager should leave a patch when travelling to and exploiting another patch yields higher return rates (Smith 1983; Bird and O'Connell 2006;

Winterhalder and Kennett 2006; Lupo 2007). A central assumption of the model is that the optimal time allocation to any patch is a function of the average return rates for all utilized patches. In general, when resources are abundant or when patches are in close proximity to one another, foragers should not spend much time in a patch. However,

57 when food availability declines or when travel time between patches increases, it becomes more profitable to increase time residency (Smith 1983; Stephens and Krebs

1986). Ultimately, patches will be ignored unless their marginal return rate equals to or is greater than the average rate for the utilized set of patches (Smith 1983; Lupo 2007).

The previous models assume that foragers consume prey at the point of capture.

However, foragers may preferentially transport resources to a central place for consumption or provisioning (Orians and Pearson 1979; Cannon 2003). This issue is addressed by central place foraging models.

4.1.3 Central place forager prey choice model

Central place foraging (CPF) models (e.g., Orians and Pearson 1979; Schoener

1979) are designed to examine how travel and transport costs influence foraging decisions regarding prey choice, load size, patch selection, and patch time allocation (Lupo 2007).

Following the assumption that the goal of foragers is to maximize their net rate of energy delivery to a central place, these models seek to predict which prey types foragers should transport and how their decisions vary as a function of the distance from a central place

(Orians and Pearson 1979). However, most CPF models do not take into account how the processing of resources at their location of acquisition can increase the utility of a load.

Small prey items may not need to be processed. In contrast, when foragers capture large prey, they often need to decide which parts to bring home and which ones to leave behind so as to maximize the net delivery rate (e.g., Metcalfe and Barlow 1992; Bird and Bliege

Bird 1997; Cannon 2000, 2003; Bird and O'Connell 2006).

Cannon’s (2003) central place forager prey choice model is designed to predict foraging decisions regarding prey selection and field processing for foragers who travel

58 from and to a central place, taking into account that there is a maximum load size that can be transported. Similar to the prey choice model, the CPF prey choice model ranks prey types in decreasing order as a function of their profitability. However, unlike the former which only considers post-encounter return rates for ranking prey types, the latter also takes into consideration search, travel, and, when it applies, field-processing costs to estimate the utility of prey types (Cannon 2003). Because search costs are included in the calculation of delivery rates, a taxon ranked high when abundant near the site is considered of lower value when found in distant patches. Additionally, the CPF prey choice model does not assume that all resources are searched for simultaneously. On the contrary, Cannon (2003) indicates that central place foragers often leave on foraging trips with a specific resource in mind, even though they might come back with a different one.

Furthermore, the model does not presume that resources are clumped in space, although it is compatible with the presence of patches in the environment (Cannon 2003).

In Cannon’s model, the relationship between processing time and load utility is assumed to approximate a diminishing-returns function (Cannon 2003). When search or transport costs increase, foragers should invest more time in field-processing large-bodied prey taxa in order to maximize the net delivery of food to the central place (Cannon

2003). Consequently, the analysis of skeletal part representation may provide information on past foraging strategies. Skeletal profiles reflecting low average search times should include a wide range of parts, including low-utility portions, while higher average search times should result in assemblages heavily dominated by highly-ranked portions.

If the goal of central place foragers is to maximize the net rate of energy delivery, these foragers should select the prey types that provide the highest delivery rates.

Considering equivalent search, transport, and field processing times, large prey types (i.e.,

59 high-ranked prey) should be more profitable than small-sized animals (Cannon 2003).

However, if the abundance of highly profitable prey taxa in patches near the camp declines, the cost associated with their acquisition increases. In this situation, an initially high-ranked taxon may become a new, lower-ranked prey type if it must be acquired in a remote location. The CPF prey choice model predicts that, at this point, foragers may concentrate their efforts on the procurement of lower-ranked, smaller-sized resources that are abundant near the residential site (Cannon 2003). However, if the search and transport costs of the small taxa increase as well near the site, foragers should increase their net delivery rate by focusing on high-ranked resources located in distant patches. This prediction of the CPF prey choice model implies that an increase in the abundance of large prey types relative to smaller ones, combined with an increase in field-processing intensity, may signal decreased rather than increased foraging efficiency (Cannon 2003).

4.2 Archaeological applications of foraging theory

4.2.1 Use of foraging models in this study

Foraging models are commonly used in archaeology to make predictions about foraging behaviours and to test hypotheses about resource depression and foraging intensification. The models are most frequently used to assess changes in foraging efficiency on the long term (Bird and O'Connell 2006; Lupo 2007). However, such application may be limited at Pacbitun. Although the faunal material was recovered from two distinct occupations (i.e., early Middle Preclassic and late Middle Preclassic), small sample size may require the assemblages to be combined, which prevents temporal comparisons between the two phases of the Middle Preclassic period. Given that the

Maya were sedentary farmers, the CPF prey choice model was selected to investigate the

60 exploitation of animal resources at Pacbitun. In accordance with Cannon’s model, the diet breadth at Pacbitun will be characterized through the use of prey ranking and abundance indices (see below). Foraging efficiency and acquisition of resources will also be examined through a consideration of transport decisions.

4.2.2 Prey rankings

Applications of foraging theory to archaeology must start with the ranking of prey types. Traditionally, prey body size has been used as a proxy for estimating the net return rate of resources (e.g., Broughton 1994; Cannon 2003; Broughton et al. 2011). In these studies, the relationship between body size and profitability is assumed to be curvilinear.

Very small prey types, such as mice, and very large animals, such as whales, are not considered as profitable as prey of intermediate sizes because a considerable amount of energy must be invested in handling them. Between the two extremes, it is generally considered that profitability increases with body size; large prey animals (e.g., deer and tapir) are ranked higher than smaller animals such as rabbits or agoutis (Broughton 1994).

Hunting technology may, however, impact return rates. For instance, the profitability of some small animals, such as rabbits, may be increased if these are collected en masse

(Jones 2006); this situation would be in contradiction with the body size because it alters the relationship between prey body size and energetic return rates. Additionally, gregarious species may be considered more profitable than solitary ones because many individuals may be handled in one foraging bout. In the Pacbitun assemblages, such animals would be the peccary and coati (see section 4.3).

The use of the prey body size as a proxy for prey profitability also fails to take into account the impact of prey mobility on return rates. Indeed, high prey velocity should

61 result in decreasing probability of capture (Stiner et al. 2000; Bird et al. 2009; Bird et al.

2012). In order to examine this issue, Morin (2012b) compared the body mass and maximum running speed of a broad range of mammals. He concluded that the body size rule works best for mammalian taxa within the size range of 50–700 kg. Unfortunately, most mammals inhabiting tropical forests, with the exception of the tapir and jaguar, weigh less than 50 kg. For these last species, rankings based on body size can be confounded by differences in prey mobility and predatory defense mechanisms (Stiner et al. 2000; Bird et al. 2009; Bird et al. 2012). Sessile and slow-moving animals using carapaces or quills (e.g., molluscs, tortoise, porcupine) to deter predators are generally associated with lower pursuit costs and higher chances of capture than animals of similar body weight (e.g., agouti, rabbit) that escape at great speed (Stiner et al. 2000; Bird et al.

2009; Morin 2012b). This situation result in a violation of the body size rule if a small- sized animal was to be found to be more profitable than a larger one. This seems to be the case in Australia where Bird and colleagues (2009; 2012) suggest that prey mobility is often a better criterion for estimating the likelihood of capturing small prey items (<25 kg) than body mass.

Morin (2012b) suggests that, when prey types are well-separated from one another in term of body size, overlap in return rates and, therefore, rank inversions are less likely.

To test this hypothesis, he compared the net return rates and body masses of faunas from different latitudinal environments. He observed that the body size rule is stronger in cooler habitats where faunal communities are comprised of several large-sized taxa of variable body weights. This implies that the body size rule may not be the most robust measure of profitability for tropical environments where faunas are generally characterized by a narrow range of similarly-sized animals. Concerning this issue,

Broughton and colleagues (2011) argue that when two taxa are well-separated in body size and are considered of similar mobility (e.g., “fast” prey such as hare vs. deer), the larger taxon should provide a higher return rate than the small prey type in accordance with the body size rule (Broughton et al. 2011). However, prey mobility may have a stronger impact on prey rankings and cause rank inversions when the faunal data set only includes a very narrow range of small-bodied prey (<30 kg).

The construction of prey rankings at Pacbitun was established using body mass as a proxy for net return rates. However, as a result of the above concerns, interpretations of the results at the site will need to consider the issue of mobility, en masse collection, and gregariousness of certain species. Body weights of mammalian taxa were obtained from

Emmons (1997) and Reid (2009) (Table 4.1 and Figure 4.1). Although carnivores were likely procured by the Maya for raw materials (e.g., pelt, teeth, and long bone shafts) or ritual purposes (Pohl 1983; Hopkins 1992; Emery 2010), they could also have been consumed. Therefore, they were included in the rankings for comparisons with other taxa.

The tapir would have been the highest-ranked species by a considerable margin, followed by a group of highly-ranked prey taxa comprised of large felines (i.e., jaguar and puma) and artiodactyls (i.e., white-tailed deer, white-lipped peccary, red brocket deer, and collared peccary). The ocelot, paca, margay, nine-banded armadillo, coati, and agouti form a second group of intermediate sizes (3–10 kg). Four animals, the opossum, rabbit, pocket gopher, and long-tailed weasel, occupy the lowest positions on the ranking scale

(<3 kg). Turtles were not included in prey ranking because, as slow-moving animals, they possibly were always included in the optimal diet despite their small size (Morin 2012b), in violation with the body size rule. Small sample size and limited taxonomic identification made the ranking of fish and bird species impracticable in the present case.

Table 4.1 Data and references for body mass of mammalian taxa at Pacbitun.

Common Name Mass (in kg) 1 Source Comparative specimens Tapir 180.0 Reid (2009) Jaguar 65.0 ” Puma 44.5 ” White-tailed deer 34.0 ” White-lipped peccary 33.5 ” Red brocket deer 22.0 ” Collared peccary 19.0 ” Ocelot 10.8 ” Paca 8.5 ” Nine-banded armadillo 5.0 ” White-nosed coati 4.6 ” Margay 3.8 ” Agouti 3.5 ” Opossums 1.5 ” (Didelphis marsupialis) Rabbits 1.0 ” (Sylvilagus brasiliensis) Pocket gophers 0.8 Emmons (1997) (Orthogeomys hispidus) Long-tailed weasel 0.5 ”

1 When a range of weights was available or when values were given separately for males and females, midpoints were used to calculate body mass.

Tapir Jaguar Puma White-tailed deer White-lipped peccary Red brocket deer Collared peccary Ocelot Paca Nine-banded armadillo White-nosed coati Margay Agouti Opossums Rabbits Pocket gophers Long-tailed weasel 0 20 40 60 80 100 120 140 160 180 Mass (in kg)

Figure 4.1 Ranking of mammals at Pacbitun according to body mass. Data from Table 4.1.

Animal resources may have been procured by the Maya not only for their meat, but also for their fat content. In the Maya subarea, certain animals of intermediate size, such as the agouti, paca, and armadillo, are lean during the dry season, but during the wet season, they can put on layers of fat greater than 2.5 cm in thickness (Hill et al. 1984).

This proportion of fat to body weight is extremely high compared to that of the white- tailed deer, whose fat content fluctuates from 2% during the dry season to 10% during the wet season (Hill et al. 1984). This difference in fat content may cause rank inversions if the Maya would have targeted small fatty animals, particularly during the wet season.

Unfortunately, it is not possible to test this hypothesis at Pacbitun because of a lack of data. Percentage of fat relative to body weight is difficult to estimate and is not reported for a large number of taxa (Morin 2012b). For instance, data on fat percentage to body weight was only found for four mammals in the Pacbitun assemblages, which is not sufficient for ranking prey according to this criterion.

4.2.3 Abundance indices

Abundance indices are often used as proxy estimates to interpret variations in the faunal record. Abundance indices generally compare a group of high-ranked prey to a group of lower-ranked taxa, taking the form of Ʃ large taxa / (Ʃ small taxa + Ʃ large taxa)

(Morin 2012b; Ugan and Simms 2012). These subsets may be established for prey of similar mobility but of different sizes (e.g., leporids versus artiodactyls in Cannon 2003;

Broughton et al. 2011) or, inversely, for prey of similar body size, but of different maximum velocity (e.g., tortoises versus hares in Stiner et al. 2000). Grouping prey types into high- and low-return categories limits the problem of rank inversions and strengthens

65 the relationship between return rates and body size. However, the use of these indices is appropriate only if the two subsets are composed of taxa with clear differences in ranking.

Errors may arise if animals of similar profitability (e.g., paca vs. agouti) are compared.

Abundance indices should also exclude small-bodied taxa that are likely to be collected en masse (Morin 2012b). The abundance indices used in this study are presented in greater details in Chapter 7.

4.3 Animal ecology and behavior

This section provides a summary of the ecology and behavior of animals recovered in Middle Preclassic deposits at Pacbitun. It serves as a basis for interpreting taxonomic representation and habitat use in subsequent chapters. It should be noted that reliable information on many Neotropical animals is scarce. Often, little is known about the seasonality and reproduction cycle of these animals because of limited research in the lowland Maya region (Ojasti 1996; Emmons 1997; Emery and Brown 2012).

The white-tailed deer is one of the largest herbivores found in the tropics. It is ecologically plastic and can live in a wide variety of habitats, such as tropical savannas, and lightly wooded or swampy areas (Ojasti 1996; Emmons 1997; Reid 2009), but is not a rainforest animal (Ojasti 1996; Emmons 1997; Reid 2009). It thrives in mosaic environments and most often resides at the edge of open and covered landscape (Ojasti

1996; Geist 1998). In Belize, the white-tailed deer is often seen in second growth forests and thickets, forest edges, pine savannas, and sometimes in milpas (Méndez 1984). It is tolerant of altered habitats, adapting quickly to changing environments (Ojasti 1996).

Active day and night, the white-tailed deer is most often spotted at dawn or dusk when it ventures in open areas to feed (Ojasti 1996; Reid 2009). The white-tailed deer is a

66 browser and selective feeder that concentrates on plants that are easily digested and feeds predominantly on leaves, fruit, nuts, seeds, and legumes (Méndez 1984; Brown 1994;

Ojasti 1996; Emmons 1997; Reid 2009). Highly fibrous plants, such as grasses, contribute little to the diet, but may be consumed when herbaceous forage is unavailable (Verme and

Ullrey 1984; Kroll 1994; Ojasti 1996). The white-tailed deer may also feed on crops, such as maize, sorghum, and beans (Ojasti 1996).

In the tropics, the white-tailed deer lives in small groups of two to six individuals, but it may also be solitary (Ojasti 1996; Emmons 1997; Reid 2009). Family herds occupy permanent small home ranges (Ojasti 1996). Although reproduction of this animal is synchronous in the north, breeding and fawning seasons extend for several months in

Central America (Ojasti 1996; Geist 1998). In fact, breeding can take place at any time of the year, but it generally peaks in certain seasons so that fawning does not occur during times of adverse weather (Geist 1994; Hirth 1994; Geist 1998). As a result, peaks in birthing are generally synchronous within a given locality (Hirth 1994; Jacobson 1994;

Ojasti 1996). For instance, fawns are born from April to June in the Yucatán and southern

Mexico, from January to June in Honduras, and from May to July in Costa Rica (Ojasti

1996). Females generally give birth to one fawn per year (Ojasti 1996). Tropical white- tailed bucks bear branched antlers, but these are smaller in comparison to northern whitetails. Because the antler cycle is irregular in the tropics, males may be encountered in any stage of antler growth (Geist 1998). Antlers are shed every year (Méndez 1984;

Geist 1998; Reid 2009).

The red brocket deer is a small tropical ungulate adapted to live in forested environments (Reid 2009). Its small size, short antlers, and rounded back allow it to move through dense vegetation (Ojasti 1996; Emmons 1997). As a result, it is not surprising

67 that this animal favors closed, mature forests, although it may occasionally visit small clearings (Emmons 1997; Reid 2009) and cultivated areas (Ojasti 1996; Emmons 1997).

This animal primarily feeds on fruit and flowers found in the underbrush, but may also browse in forest clearings (Ojasti 1996; Emmons 1997). The red brocket deer is a solitary and territorial animal; it occupies the same home range year after year (Ojasti 1996).

Bucks bear short, spiked antlers (Emmons 1997; Geist 1998; Reid 2009) which may be retained for more than a year but can be shed at any time (Geist 1998). Similar to most tropical mammals, reproduction in brocket deer may take place year-round (Ojasti 1996;

Geist 1998). Peaks in birthing occur at specific times of the year in different regions.

Fawns are born between September and April in Surinam, whereas the calving season extends from April to August in Chiapas, Mexico (Ojasti 1996). The red brocket deer normally gives birth to a single young (Ojasti 1996; Reid 2009).

Morphologically similar to a pig, the white-lipped peccary is a medium-sized ungulate (Mayer and Wetzel 1987). This gregarious animal lives in large herds formed of

40 to 200 individuals of both sexes (Donkin 1985; Mayer and Wetzel 1987; Ojasti 1996;

Emmons 1997; Reid 2009). Its preferred habitats are virgin or near-pristine humid tropical forests (Mayer and Wetzel 1987; Emmons 1997; Reid 2009). It generally avoids disturbed habitats and the proximity of humans (Ojasti 1996). This species is considered nomadic; herds travel long distances throughout the tropical forest and do not stay more than a day or two in a particular area (Mayer and Wetzel 1987; Ojasti 1996; Emmons

1997; Reid 2009). It subsists on a mix of fruit, leaves, seeds, roots, and small vertebrates and invertebrates (Donkin 1985; Mayer and Wetzel 1987), and may sometimes exploit crops, such as maize, sweet potatoes, and manioc (Mayer and Wetzel 1987; Reid 2009). It

68 is believed to breed year-round, with females typically giving birth to twins (Mayer and

Wetzel 1987; Ojasti 1996; Reid 2009).

The collared peccary is smaller than the white-lipped peccary (Ojasti 1996; Reid

2009). It lives in herds composed of about 15 adults and their young (Donkin 1985;

Emmons 1997; Reid 2009). The composition of the group is loose and males may occasionally leave the herd to forage alone (Ojasti 1996; Emmons 1997). The collared peccary occupies permanent home ranges (Ojasti 1996; Reid 2009), but is more versatile than the white-lipped peccary. This species can live in both mature and secondary forests and adapts well to disturbed areas (Donkin 1985; Ojasti 1996; Reid 2009). Its diet consists of fruit, seeds, roots, grasses, invertebrates, and small vertebrates (Donkin 1985;

Ojasti 1996; Emmons 1997; Reid 2009). This species commonly raids crops (Ojasti 1996;

Reid 2009). Like other ungulates in Central America, the collared peccary can breed year- round. The birthing season lasts from July to August in the southern United States and from March to April in Venezuela. Two birth peaks were observed in southern Mexico, the first extending from January to February, the second from September to October

(Ojasti 1996). Females give birth to two offspring on average (Reid 2009).

The Baird’s tapir is a large perissodactyl. This animal is normally solitary, but family groups composed of a cow and her offspring are sometimes observed (Emmons

1997; Reid 2009). The tapir favors waterside habitats with dense, low vegetation.

Nonetheless, it is ecologically plastic and travels extensively throughout the forest to forage (Emmons 1997; Reid 2009). Being a shy animal, the tapir tends to avoid human settlements, but it may occasionally be spotted raiding crops (Reid 2009).

The nine-banded armadillo occupies a wide range of primary and secondary habitats, but is most often seen in thickets and areas of dense vegetation (Emmons 1997;

Reid 2009). Sedentary, the animal dens in large burrows during the day and feeds at night on insects and small invertebrates. It may occasionally eat berries and other plants

(McBee and Baker 1982; Ojasti 1996; Emmons 1997; Reid 2009). Individuals are solitary, but show no tendency toward territoriality (McBee and Baker 1982; Reid 2009).

The periodicity and frequency of armadillo reproduction in Central America is unknown.

It is believed that the mating season coincides with the onset of rains from May to August

(McBee and Baker 1982; Ojasti 1996).

The paca is a large rodent that occupies a wide ecological range of forested environments, from mature and secondary forests to riverine habitats and wetlands (Pérez

1992; Ojasti 1996; Emmons 1997; Reid 2009). It may also occupy wooded areas near agricultural zones (Ojasti 1996; Reid 2009). This animal is mainly active at night when it leaves its underground burrow to forage on fruit, seeds, and young plants (Pérez 1992;

Ojasti 1996; Reid 2009). Living in monogamous pairs, the paca occupies an exclusive territory (Ojasti 1996; Emmons 1997). Females generally give birth to a single young

(Pérez 1992; Reid 2009). They may have one or two litters per year (Pérez 1992; Ojasti

1996).

The agouti is another large rodent inhabiting the tropical forests of Central

America. Smaller than the paca, the agouti thrives in forested habitats. Because it is fairly tolerant of modified environments, it can also be found near cultivated lands (Ojasti 1996;

Emmons 1997; Reid 2009). As a forest-dweller, this small mammal feeds on a wide variety of fruit and seeds, occasionally supplementing its diet with fungi, sprouts, and crops (Ojasti 1996; Emmons 1997; Reid 2009). Agoutis are most active in the morning or the late afternoon (Ojasti 1996; Reid 2009). A breeding pair of agoutis usually occupies a

70 territory of two to three hectares (Ojasti 1996; Reid 2009). They can breed year-round, with two litters a year as a rule (Ojasti 1996; Reid 2009).

Pocket gophers are large, rat-like rodents that live in small isolated communities.

These animals spend most of their lives underground where they build deep, complex burrow systems (Emmons 1997). They prefer open habitats with well-drained soils, such as agricultural clearings or hilly forested areas. They generally feed on plant parts that grow underground, such as tubers and stems, but they can also pull down aboveground plants from below (Emmons 1997).

Two types of rabbits can be found in Central America: the tapiti and the cottontail.

Both species are browsers and grazers (Chapman et al. 1980; Emmons 1997). The tapiti is the only type of rabbit that truly resides in mature tropical forests (Emmons 1997).

Mainly nocturnal, it commonly forages in habitat edges and agricultural fields, finding shelter in thickets of dense vegetation (Emmons 1997; Reid 2009). In Chiapas, Mexico, this species breeds year-round (Reid 2009). The cottontail prefers to live in open areas, such as savannas, grasslands, and agricultural fields (Reid 2009). Although this animal is solitary, it is not territorial (Chapman et al. 1980; Reid 2009). Females may have five to seven litters per year (Reid 2009).

The white-nosed coati preferentially lives in wooded habitats, but it is occasionally seen near savannas or agricultural fields (Gompper 1995; Emmons 1997;

Reid 2009). It spends most of its time foraging on the ground, but it may climb in the canopy to forage fruit trees or rest (Gompper 1995; Reid 2009). Adult females and their offspring travel in groups of about 20 individuals, whereas males are usually solitary

(Gompper 1995; Emmons 1997; Reid 2009). This animal is omnivorous and feeds on fruit, invertebrates, and small vertebrates (Gompper 1995; Emmons 1997; Reid 2009). In

71 the few populations surveyed, coatis breed synchronously once per year, although the timing of the breeding season differs between regions (Gompper 1995).

The long-tailed weasel efficiently occupies a wide range of environments. It can inhabit forested locales but it usually prefers open areas and agricultural lands (Emmons

1997; Reid 2009). This animal is a generalist predator (Sheffield and Thomas 1997). In tropical settings, it preys on small mammals (less than 50 g), rabbits, pocket gophers, birds and their eggs (Sheffield and Thomas 1997; Reid 2009). Out of the den, it is active day and night; most of the time is spent foraging and feeding (Sheffield and Thomas

1997). Nests are habitually found in burrows made by other animals or under rocks

(Sheffield and Thomas 1997; Reid 2009). This animal is solitary, but a male and a female may remain together during the non-breeding season (Sheffield and Thomas 1997).

The Virginia opossum is a small, highly opportunistic animal. It can live in many types of forested and open habitats and is generally found in the wetter areas of its range, near streams and swamps. It is known to frequently forage in human refuse piles and, as a result, is commonly found near human settlements (McManus 1974; Emmons 1997; Reid

2009). The diet of this omnivorous animal consists of insects and other invertebrates, small vertebrates, carrion, fruit, and other plant matter (Emmons 1997). Mainly active at night, the Virginia opossum spends most of its time travelling and feeding on the ground although it may climb readily (Emmons 1997; Reid 2009). It is known to “play dead” when disturbed or threatened (Emmons 1997; Reid 2009).

The common opossum is similar in its habits to the Virginia opossum. It mainly lives on the ground, but may climb in trees to feed or escape danger (Emmons 1997; Reid

2009). This animal focuses on the exploitation of small animals (e.g., insects, worms, and small vertebrates), although regularly forages for fruit and other plant matter (Emmons

1997; Reid 2009). In addition, it frequently takes advantage of the presence of garbage dumps in rural areas (Reid 2009). At night, it may travel one to three kilometers in search for food, but it remains in well-defined home ranges (Reid 2009). This animal thrives in secondary forests and disturbed areas, although it is also found in mature, humid forests

(Emmons 1997; Reid 2009).

All felines found in the Maya subarea occupy the rainforests. Solitary hunters, they handle almost anything they encounter, including mammals, snakes, birds, turtles, caimans, fish, and even large insects (Emmons 1997). All species are believed to breed year-round in the tropics, with litter size varying from one to four kittens (Tewes and

Schmidly 1987; Emmons 1997; Reid 2009). All species are mainly active nocturnally

(Tewes and Schmidly 1987; Emmons 1997).

The ocelot is a small feline that targets a variety of small- and medium-sized prey

(e.g., rodents, birds, snakes, lizards, and other small vertebrates). It spends most of its time foraging on the ground and seldom climbs in trees (Tewes and Schmidly 1987;

Emmons 1997; Reid 2009). The ocelot may occupy many different habitats (e.g., rainforest, tropical deciduous forest, secondary forest, and scrubs), as long as they provide sufficient cover (Tewes and Schmidly 1987; Emmons 1997; Reid 2009). It may also live in disturbed areas and near human settlements (Emmons 1997; Reid 2009).

Slightly larger than a house cat, the margay is another small feline inhabiting tropical rainforests (Tewes and Schmidly 1987; Emmons 1997). This species is the most arboreal cat in the Americas. It hunts and rests in trees, but travelling is done on the ground (Emmons 1997; Reid 2009). As a consequence of its arboreal habits, the margay is found almost exclusively in mature and secondary forests and does not adapt well to human disturbance (Tewes and Schmidly 1987; Emmons 1997; Reid 2009). Small

73 mammals, birds, and reptiles form the bulk of its diet (Emmons 1997; Reid 2009). Strictly nocturnal, the margay spends daylight hours hidden among dense trees (Tewes and

Schmidly 1987).

Two large cats inhabit the tropical forests of the Maya subarea. The puma is a solitary animal, usually wary of humans (Emmons 1997; Reid 2009). This animal is highly adaptable and can be found in a variety of habitats, such as rainforests, scrubs, and mountains (Emmons 1997; Reid 2009). The jaguar is the largest cat in the Americas. It favors undisturbed forested habitats and waterside areas (Tewes and Schmidly 1987;

Emmons 1997; Reid 2009). Both species predominantly prey upon medium- and large- sized mammals, such as deer, peccary, agouti, and paca. Depending on local availability, they may also feed on turtles, caimans, birds, fish, and smaller mammals (Tewes and

Schmidly 1987; Emmons 1997; Reid 2009). Jaguars and pumas are sedentary animals that presumably occupy defined home ranges (Tewes and Schmidly 1987; Emmons

1997).

Both the black and green iguanas live in tropical moist and dry forests, although the black variety may also be found in open savannas (Campbell 1998). The green iguana is arboreal. It spends most of its time in trees and is normally found near permanent sources of water (e.g., rivers, streams, and lakes). In contrast, the black iguana lives closer to the ground and rests in underground burrows or hollow logs (Campbell 1998). Both species are primarily herbivorous, feeding on leaves, flowers, and fruit. It is reported that the black iguana can eat small birds, rodents, and reptiles. Females lay eggs during the dry season, from February to June (Campbell 1998).

Mud and musk turtles (Kinosternidae) are found in shallow backwaters of streams and lakes, marshes, swamps, and aguadas. Generally, these turtles are only active

74 seasonally, because they bury themselves into the mud during the dry season (Campbell

1998). The northern giant musk turtle (Staurotypus triporcatus) is the only species living near large, permanent bodies of water and active year-round. These turtles are usually active at night. They are poor swimmers and forage at the bottom of ponds and lakes.

They do not need to emerge from the water to bask (Campbell 1998). Their diet is composed of a mix of aquatic invertebrates, such as insects, shrimps, molluscs and crabs, vegetable matter, tadpoles, and carrion (Campbell 1998).

These observations about the ecology and behavior of the most prevalent species recovered at Pacbitun are central to an interpretation of the subsistence behaviors of the

Middle Preclassic Maya. However, before any interpretation can take place, the quantitative and taphonomic methods used in the analysis of the faunal remains need to be examined. They are the focus of the following chapter.

CHAPTER 5: METHODOLOGY

In order to interpret subsistence strategies at Pacbitun, it is necessary to evaluate the integrity of the study sample. Indeed, multiple pre- and post-depositional processes may have altered faunal assemblages before and after their recovery by the archaeologist.

Consequently, this chapter presents the methods used to assess taphonomic signatures on the Pacbitun assemblages, along with a description of the quantification methods used in the analysis of taxonomic composition and skeletal part representation. The chapter also considers why the chosen methods are appropriate for studying faunal assemblages recovered in tropical environments.

5.1 Definitions and identification procedures

Several important concepts used throughout this analysis must be defined prior to any discussion regarding the faunal assemblages investigated in this study. A “skeletal element” refers to a single complete anatomical unit, such as a humerus or a molar. A

“specimen” is described as any skeletal remain anatomically complete or a fragment thereof, such as a proximal humerus or a molar fragment. The implication is that all skeletal elements are specimens, but not all specimens are skeletal elements (Grayson

1984:16; Lyman 2008:5).

The identification of faunal remains was carried using comparative collections from the Archaeozoology Laboratory at Trent University and the Department of

Vertebrate Paleontology at the Royal Ontario Museum. Manuals by Olsen (1982), Barone

(1986), and Gilbert (1993) were also consulted. Vertebrate remains were identified to the highest taxonomic level possible. All identified specimens were examined using a 10X magnification lens to observe surface modifications.

5.2 Quantification methods

The Number of Identified Specimens (NISP) is a fundamental measure of faunal quantification (Grayson 1984; Lyman 2008; Reitz and Wing 2008). It is defined as the number of skeletal elements and fragments identified to taxon. Specimens counted in

NISP are generally identified to the skeletal element and to at least the genus level

(Lyman 2008:27). However, in this study, NISP values also include bird, reptile, and fish specimens identified to the class level, as these taxa could not always be identified as precisely as mammalian specimens. Isolated teeth were counted as separate elements if they could not be associated with another cranial element. If they were still in articulation with the maxilla or mandible, they were considered part of the cranial element and were not added to the NISP. When mammal bones could not be identified to family or genus, they were attributed to broader taxonomic groups (Table 5.1), following the body size classes defined by Savage (1971) and Emery (2007a).

Table 5.1 Taxonomic groups based on body size.

Small mammal Medium mammal Large mammal Gopher Domestic dog White-tailed deer Mouse Armadillo Brocket deer Squirrel Opossum Tapir Agouti Jaguar Paca Puma Small felines Peccary Rabbits Raccoons

The Minimum Number of Elements (MNE) corresponds to the minimum number of skeletal portions necessary to account for all the specimens representing a particular element (Bunn and Kroll 1986:434–435; Lyman 1994:102). MNE estimates were determined by manually counting, for each skeletal element, the number of portions with

77 overlapping landmarks (Todd and Rapson 1988) or zones (Morlan 1994) for proximal and distal ends, as well as for shafts. Derived from MNE, the Minimum Number of

Individuals (MNI) is a measure of abundance consisting of the smallest number of individuals necessary to account for all of the specimens of a particular taxon in an assemblage (Klein and Cruz-Uribe 1984; Lyman 2008). MNI values were calculated on the basis of the most common anatomical element for each taxon.

For both MNE and MNI, age, sex, and specimen size were not taken into account, because the use of these characteristics tends to inflate the representation of diagnostic specimens, such as teeth and epiphyses, in comparison to less diagnostic specimens, for instance, long bone shaft and rib fragments (Morin 2012b:68). Only stratigraphic provenience (early Middle Preclassic versus late Middle Preclassic deposits) was considered in the calculation of MNE and MNI estimates. MNE was used to measure both the frequencies of skeletal parts and the abundance of distinct individuals.

Derived from MNE as well, the Minimal Animal Units (MAU) is used to describe skeletal part frequencies (Lyman 2008). MAU values were determined by dividing MNE estimates for each element by the number of times this element is represented in a complete skeleton. The values were then standardized (%MAU) by dividing all MAU values by the greatest MAU value in the assemblage (Binford 1984:50). This procedure reduces much of the variation in MNE values caused by the differential frequency of skeletal elements in a complete skeleton (Lyman 2008:233–237).

A similar protocol was adopted for the use of NISP in the analysis of skeletal part frequencies. NISP counts for each skeletal element were divided by the number of times this element occurs in a particular taxon, resulting in a Normed NISP (NNISP) (Grayson and Frey 2004; Grayson and Delpech 2008). For example, NNISP values for long bones

78 were obtained by dividing the NISP counts by two. The same procedure was applied to cranial elements, as crania and mandibles are represented in the assemblages by isolated left and right fragments. NNISP counts were also standardized (%NNISP) to the abundance of the most common element for each taxon.

5.2.1 Strengths and weaknesses of NISP, MNE, and MNI

The main advantage of NISP lies in its simplicity. NISP is a direct tally of identified specimens. As new specimens are identified and added to the analysis, the researcher does not have to recalculate the totals. NISP is also easily replicable. Two different observers, assuming comparable skills in identification and access to the same collections, should obtain similar NISPs for the same faunal assemblage (Grayson

1984:20; Klein and Cruz-Uribe 1984:25; Lyman 2008:28). In addition, because it is cumulative, NISP does not suffer from problems of aggregation between units or layers

(Grayson 1984). One specimen is always considered to be one specimen no matter where it was found or how it is aggregated. Finally, as different analysts would likely tally NISP more or less the same way, NISP has the advantage of facilitating comparisons of assemblages from different sites. This is particularly true for Maya archaeozoology given that the majority of lowland Maya faunal analyses are based on NISP counts (Emery

1997:90).

NISP does, however, suffer from various shortcomings. First, NISP is sensitive to inter-taxonomic and intra-taxonomic variation. Indeed, different taxa do not share the same number of skeletal elements and some taxa or elements may be more easily identified than others. Some skeletal elements or taxa are also more likely to be fragmented or destroyed as a result of differential structural density (Grayson 1984:20–

25; Klein and Cruz-Uribe 1984:25; Marshall and Pilgram 1993; Ringrose 1993; Lyman

2008:29–34). Given these problems, the abundance of certain elements or taxa may be under- or over-represented relative to their initial abundance in the deposited assemblage as assessed by NISP. Differential identification and fragmentation can be mitigated by refitting (discussed in section 4.3). Refitting is defined as piecing back together specimens that belong to the same skeletal element (mechanical refit) or by reassembling together skeletal elements from a same individual (anatomical refit) (Enloe and David

1989, 1992; Hofman 1992). Differential representation caused by differences in total number of elements between species can be controlled for with the use of additional quantification methods, such as normed NISP (NNISP).

NISP is also plagued by the problem of interdependence (Grayson 1984; Lyman

2008; Reitz and Wing 2008:203), given that it does not account for the fact that multiple specimens identified to a taxon may be from a single individual. For instance, if the NISP is 50, it is not possible to determine whether those specimens represent 1 or 50 individuals. In this context, NISP may exaggerate the sample size of individuals or elements. The problem of interdependence may preclude the use of basic statistical tests as most require independent observations (Grayson 1984:23–26; Lyman 2008:36–38). A high degree of fragmentation may also amplify the problem of interdependence (Klein and Cruz-Uribe 1984:25; Marshall and Pilgram 1993). This drawback of NISP can partially be countered by conducting refit studies and by using derived quantification measures, such as the minimum number of individuals (MNI). MNI solves the problem of interdependence as it avoids counting the same individual twice. Similarly, it is not affected by the problem of inter-taxonomic variation of skeletal elements, as it is based on the most common skeletal element of a taxon in an assemblage (Lyman 2008:38–39, 44).

Although MNI was introduced to overcome the problems associated with NISP counts, several problems with this quantification method have also been identified. MNI is more difficult to tally than NISP as it is not cumulative. Moreover, there is no consensus on how MNI estimates should be derived (Klein and Cruz-Uribe 1984:26;

Marean et al. 2001; Lyman 2008:46), which may reduce comparability between assemblages. This problem can, to some extent, be controlled for if the same methods are used (Klein and Cruz-Uribe 1984:26–28; Marean et al. 2001; Lyman 2008:46–48).

Additionally, MNI counts can exaggerate the importance of rare taxa. This problem can be assessed if MNI estimates are considered together with NISP values (Klein and Cruz-

Uribe 1984:32–33; Plug and Plug 1990; Lyman 2008:46–47).

MNI values can also be affected by sample aggregation. The MNI count for an entire site will likely differ from the MNI estimates calculated for multiple distinct units of the same site, because a given skeletal part might not constitute the most abundant element across different aggregates. For instance, although the left humerus of a given taxon might be the most abundant skeletal part at a site, other elements might occur more frequently if the assemblage is divided in multiple aggregates (Grayson 1984:29–30; Plug and Plug 1990; Lyman 2008:58–63). Refitting can help overcome the problem of aggregation, as it reduces the possibility of aggregates sharing specimens from the same individuals (Lyman 2008:68–69).

The nonlinear increase of MNI with increasing sample size also constitutes a problem. Similar to NISP, MNI increases with fragmentation. However, as the intensity of fragmentation increases, it becomes increasingly difficult to identify fragments because they retain fewer and fewer diagnostic landmarks (Lyman and O'Brien 1987; Marshall and Pilgram 1993; Lyman 2008:43). At some point, the rate of increase of MNI will

81 decrease relative to the rate of increase of NISP (Marshall and Pilgram 1993) because the probability of adding a specimen of a new, unrepresented individual to MNI counts decreases concomitantly (Grayson 1984:61–64; Lyman 2008:48–50).

Another derived measure, MNE, has many of the same advantages and problems.

Like MNI, MNE is used to control for the problem of interdependence. MNE ensures that each element will not be tallied twice which helps to circumvent the problem of differential fragmentation. At first glance, MNE may represent a better unit than NISP for quantifying the abundance of skeletal parts (Lyman 2008:222). However, like MNI, MNE is influenced by strategies of aggregation and sample size (Lyman 2008:222–224).

Due to the problems of differential preservation, fragmentation, and identification,

NISP and MNI values cannot be considered as ratio-scale data (Grayson 1984:94–96).

Nonetheless, NISP and MNI can provide reliable ordinal data when taxa are well separated in terms of abundance. Indeed, the greater the difference in abundance between taxa, the less likely that specimen interdependence (for NISP) or the use of different aggregation methods (for MNI) will alter the rank order of those taxa (Grayson 1984:97–

99; Lyman 2008:73–75). The same reasoning applies to MNE estimates (Lyman

2008:226–228).

5.2.2 Use of quantification methods in this study

The small size of the Pacbitun assemblages may affect and limit the use of certain quantification methods. MNE derived counts, such as the MNI and MAU, are not, in comparison to NISP, particularly accurate when used to describe small samples (Grayson

1978; Plug and Plug 1990). This problem is also amplified by the high species diversity that characterizes the assemblages. As a result of small sample size, the results obtained

82 from statistical tests based on MNE and MAU values may not be very robust. To limit the problems associated with each quantification unit, NISP and MNE were used conjointly to measure taxonomic composition, whereas NNISP and MAU values were used to investigate skeletal part representation. The use of several quantification methods should increase the robustness of the results with respect to taxonomic composition and skeletal part representation.

5.3 Refitting

Refitting is useful because conjoining two or more fragments together can increase the identifiability of otherwise nondiagnostic fragments, particularly long bone shaft fragments (Marean and Kim 1998). Additionally, the stratigraphic integrity of an archaeological sequence can be tested through an analysis of the provenience of the refitted fragments (Hofman 1992; Morin et al. 2005). Although refitting an entire faunal assemblage is tedious and time-consuming (Marean and Kim 1998), even partial refitting of an assemblage can provide substantial information on occupation mixing (Morin et al.

2005).

Because of the small sample size of the Pacbitun assemblage, it was possible to attempt to refit all long bone shaft fragments, rib fragments, and taxonomically identified specimens. Efforts were made to refit unidentified shaft fragments with other specimens from the same unit. Identified specimens were tentatively refitted with other fragments from the same unit, as well as with all specimens belonging to the same element and taxon. This procedure was carried out across all levels. A distinction was made between dry bone and green bone given that they can yield information on site formation processes and occupation mixing (Todd and Stanford 1992).

5.4 Age and sex

The analysis of animal age and sex can produce important information on seasonality of occupation, human prey choice, and husbandry practices. Epiphyseal fusion is a common method used to estimate the age at death of particular individuals in faunal assemblages. Because bones often fused successively during the life of animals, the sequence of fusion can be used to establish coarse age classes, particularly for juvenile specimens (Reitz and Wing 2008:72). However, when all the bones of a skeleton are fused, it is seldom possible to distinguish between stages of older individuals (e.g., prime adults versus senile individuals). Therefore, epiphyseal age classes may only provide coarse information on mortality profiles. This problem may be accentuated by the fact that unfused epiphyses are more easily affected by destructive processes (e.g., carnivore gnawing, post-depositional processes) than fused ones, which can lead to biases against younger individuals (Klein and Cruz-Uribe 1984:41–43).

Fortunately, the use of teeth for estimating age frequently yields more precise information than epiphyseal fusion. First, teeth are generally more durable than epiphyses and are less likely to be damaged by post-depositional destruction (Klein and Cruz-Uribe

1984:43–44). Second, teeth erupt at a certain age and wear down more or less continuously throughout the life of an animal (Klein and Cruz-Uribe 1984:43–44; Reitz and Wing 2008:72–73). Consequently, the relative age of mammals can often be estimated by using extant sequences of dental eruption and wear (Klein and Cruz-Uribe

1984:46–55; Hillson 2005:215–222). Age for worn teeth is frequently measured with the maximum height of the crown of cheek teeth, both deciduous and permanent (Payne

1973; Hillson 2005:214–222; Reitz and Wing 2008:72–73). However, the method

84 requires complete teeth, which can limit the number of specimens suitable for constructing age profiles (Klein and Cruz-Uribe 1984:52–54). Additionally, there can be significant variation in dental eruption and wear between populations (Klein and Cruz-

Uribe 1984:52) and, in some species, between sexes (Gee et al. 2002; Hillson 2005:223).

Data is not always available for wild animals (Klein and Cruz-Uribe 1984:41–43), but both biologists (e.g., Severinghaus 1949; Gilbert and Slolt 1970; Gee et al. 2002) and archaeologists (e.g., Elder 1965; Wolverton et al. 2008) have been successful in using this method for white-tailed deer, the most common species at Pacbitun.

At Pacbitun, age determination focused on white-tailed deer. Data on epiphyseal fusion was compared to sequences recorded by Purdue (1983). Epiphyses were coded as unfused, intermediate, or fused. Bones in the intermediate category are those for which the line of fusion is still clearly visible. Age from teeth was estimated using measurements provided by Severinghaus (1949). Crown height measurements were recorded for mandibular premolars and molars. Following the approach adopted by Gee and colleagues (2002), specimens were assigned to one of three age categories: juvenile

(<1 year old), subadult (1–2 years old), and adult (>2 years old). The adult class was subsequently split into young and old adults based on dental wear. When possible, age was also determined for other species if published sequences of epiphyseal fusion or dental eruption and wear were available.

Concerning sex determination, one of the simplest methods consists of recording the presence or absence of antlers for ungulates (Klein and Cruz-Uribe 1984:39–40). Sex can also be determined based on osteometric differences between males and females.

However, this technique requires bone specimens sufficiently complete to allow adequate

85 measurements to be taken (Klein and Cruz-Uribe 1984:39–41). The absence of antlers, small sample size, and substantial fragmentation prevented sex determination at Pacbitun.

5.5 Taphonomic modifications

Anthropic, biological, chemical, and physical factors may all affect archaeological assemblages and distort the interpretation of human behaviours inferred from faunal analyses. Therefore, multiple lines of evidence are examined here in order to determine which agent(s) is(are) responsible for the formation of the faunal assemblages at Pacbitun.

5.5.1 Fractures

Although taphonomic and actualistic studies have shown that there is no simple correlation between a type of fracture and the agent that produced it (Binford 1981;

Haynes 1983b; Johnson 1985), the study of fracture morphology can help to infer the timing of bone fracturing. The importance of studying fracture morphologies on long bone fragments should be emphasized because fractures constitute constant properties of broken bones (Villa and Mahieu 1991). It may be possible to study fractures even when cortical surfaces are heavily weathered, as is frequent in tropical environments, or when cutmarks or gnawing marks have become obscured by poor surface preservation.

In this study, fractures on long bone fragments have been assigned to two types: green-bone and dry-bone fractures. Green-bone fractures tend to have a homogeneous color and show a smooth texture and spiral shape (Morlan 1980:48–49; Haynes 1983a;

Johnson 1985:176, 222). Dry-bone fractures often exhibit a rough texture in cross-section, have a more or less transverse shape and may present a different color than the outer cortical surface (Morlan 1980:48–49; Johnson 1985:176–178, 222). Only unambiguous

86 fractures were recorded following this procedure. Although this method has mainly been used with mammal remains, it can also be applied to the analysis of avian long bones

(Higgins 1999; Serjeantson 2009). Additionally, following the terminology of Villa and

Mahieu (1991), the shape of proximal and distal fractures on long bones was assigned to one of the following types: curved, v-shaped, oblique, transverse, irregular, or ragged.

Fracture edges were recorded following Villa and colleagues (2005) as fresh, slightly abraded, abraded, and very abraded, as this information can potentially help to infer about site formation processes.

5.5.2 Butchery and tool use

Butchery can be defined as all sets of human activities, such as skinning, disarticulation, and meat removal, associated with the extraction of consumable resources from an animal carcass (Lyman 1994:294–296). Cutmarks can result from such actions

(Binford 1981:46–47; Bunn 1983b), as well as from the manufacture of bone tools

(Emery 2008a). Therefore, cutmarks constitute the best indicator of human involvement in assemblage formation (Villa et al. 2004). Because they can be mistaken with marks produced by the action of carnivores or by trampling, only unambiguous cutmarks were recorded in this analysis.

Marrow constitutes an important source of fat and nutrients. Bones are generally cracked open by humans with a hammerstone-on-anvil technique, which may leave percussion marks and notches (Johnson 1985:192–194; Pickering and Egeland 2006).

Because they penetrate the thickness of the bone, notches are less easily obscured by weathering than marks left on the cortical surface of the bone. As a result, they can be used, with some limitations, to identify the agents responsible for the formation of

87 archaeological assemblages (Capaldo and Blumenschine 1994). Unfortunately, notches made by carnivores can be very similar to notches produced by humans (Bunn 1989).

Nonetheless, Capaldo and Blumenschine (1994) suggest that the morphology of notches produced by humans and carnivores can be statistically distinguished for small bovids

(less than 115 kg). In a recent study, Galán and colleagues (2009) observed considerable overlap between carnivore-induced and hammerstone-induced notches on small bovid assemblages and concluded that the pattern for small bovids is much weaker than proposed by Capaldo and Blumenschine. Castel (2004) also noticed that, in some instances, the morphology of notches produced by wolves on sheep bones can be similar to that produced by human activities. Despite this lack of agreement in the literature, all notches were recorded in this study. Because of the marginal presence of carnivores in the

Pacbitun assemblages (see Chapter 6), it is believed that notches are probably the result of anthropogenic fracturing.

The Maya manufactured a wide range of artifacts (e.g., needles, scrapers, fish- hooks, beads, pins, and musical instruments) from bone, tooth, and antler (Hamblin 1984;

Pohl 1990; Moholy-Nagy 1994:110; Emery 2008a, 2009). Bone artifacts recovered from

Maya sites, such as Tikal (Moholy-Nagy 1994) and Dos Pilas (Emery 2008a, 2009), indicate that these were generally made of mammalian bones. The Maya preferentially selected the straightest and strongest bones for this purpose, in particular the tibia, femur, and metapodial (Hamblin 1984; Emery 2008a, 2009). Bones from white-tailed deer was the preferred raw material, but other medium and large mammals (felids, peccary, dog, and brocket deer) were also exploited (Moholy-Nagy 1994:107; Emery 2008a, 2009).

Fragile cranial fragments and scapulae might also have been selected for the production of disks and other flat artifacts (Moholy-Nagy 1994:107; Emery 2008a, 2009). In some

88 cases, the edges of broken bones were used as tools without further shaping. These expedient tools can be identified by the polish visible on the bone surface (Pohl

1990:158). One artefact made from a mammal long bone was identified in the sample.

Bone debris resulting from artifact production may also be present, but were not identified during the analysis.

5.5.3 Carnivore ravaging

Carnivores can be important agents of accumulation and transformation of faunal assemblages. Unfortunately, studies focusing on the action of carnivores in tropical environments are rare. The literature, however, abounds with actualistic and archaeological studies in other areas of the world (e.g., Binford 1981; Marean and

Spencer 1991; Stiner 1994). These provide considerable insight on how to recognize the signature of carnivore agents in faunal assemblages.

The most destructive agent at Maya sites is likely to have been the domestic dog

(Stanchly 2004). As companions, dogs possibly scavenged household waste or were fed butchery scraps (Pohl 1990; van der Merwe et al. 2000; Stanchly 2004). Therefore, they could have both accumulated and transformed the faunal assemblages at Pacbitun. Dogs can cause extensive damage to animal bones. They often destroy the cancellous epiphyses, gnawing the ends into the shaft to extract fat. Relatively complete shafts or

“bone cylinders” may remain from this process (Binford 1981; Johnson 1985:191–192;

Cruz-Uribe 1991; Castel 2004). In contrast, humans tend to break bones along the mid- shaft, which results in a high proportion of fragmented bones or “splinters” (Bunn 1983a;

Pickering and Egeland 2006). Therefore, long bone shaft fragmentation (see section

4.5.4) is an important aspect to consider when determining the impact of carnivores on a

89 faunal assemblage. It should be noted that dogs can also fracture bones through static loading, producing notches and pits (Johnson 1985:192; Capaldo and Blumenschine

1994; Castel 2004). In dog experiments, punctures and furrows occur frequently on the epiphyses, while scoring and pitting are more common on the diaphysis (Binford

1981:44–49). Dogs also tend to swallow bone fragments or small bones while eating.

These bones may be chemically eroded to varying degrees during digestion (Andrews and

Evans 1983).

Pumas and jaguars also accumulate and modify faunal assemblages, but not as actively as other carnivores (Seymour 1989; Martín and Borrero 1997). Large cats rarely cause heavy gnawing damage, because they do not tend to consume grease and marrow contained in bones. However, they can leave few, relatively deep and large punctures and furrows on the epiphyses (Haynes 1983b; Martín and Borrero 1997; Montalvo et al. 2007;

Mondini and Muñoz 2008; Muñoz et al. 2008). Smaller felids, including the ocelot, margay, and jaguarundi, may also significantly damage small mammal (<5 kg) bones

(Álvarez et al. 2012), but are less likely to cause important damage to large carcasses.

Although felids do not tend to come near human settlements and rarely scavenge on carcasses (Seymour 1989), they nonetheless could have accumulated or modified the faunal assemblages at Pacbitun.

Other animals present in the habitats surrounding Pacbitun can potentially accumulate or alter faunal assemblages. Procyonids (e.g., coati, raccoon, and kinkajou) are omnivorous. They feed mainly on fruits and invertebrates, and occasionally on small rodents and birds (Lotze and Anderson 1979; Ford and Hoffmann 1988; Gompper 1995,

1996). Mustelids, including the long-tailed weasel and tayra (Eira barbara), are carnivorous. They consume a variety of insects and small vertebrates, including small

90 rodents, rabbits, iguanas, and birds of small to medium size (Sheffield and Thomas 1997;

Presley 2000). Therefore, despite their small size, mustelids and procyonids may prey on the same small taxa as humans do. Some species may live near human habitations, taking advantage of food found in gardens, orchards, and agricultural fields (Ford and Hoffmann

1988; Gompper 1995, 1996; Presley 2000). These carnivores probably have little impact on the bones of large animals, because their carnassial teeth are poorly developed compared to larger carnivores (Ford and Hoffmann 1988; Gompper 1995). However, they are known to accumulate small-sized vertebrates, particularly in dens and burrows

(Andrews and Evans 1983). Lastly, rodents, such as the agouti and paca, also transport and gnaw bones. Rodent gnaw marks can easily be identified because of the distinctive furrows they produce with their incisors (Binford 1981; Johnson 1985:180).

Following Binford (1981:44–51), carnivore gnaw marks were classified in this study as tooth pits, punctures, crenulated edges, scoring, furrows, scooping, and traces of digestion. The extent of carnivore gnawing was also recorded as marginal, limited to one section, or extensive (Morin 2012b:73). Because both surface preservation and bone fragmentation can affect the observation of marks on bones, data on fragment size, completeness and circumference of long bone shafts is also required to understand animal activity.

5.5.4 Fragmentation

The study of bone fragmentation can reveal the extent of anthropic and animal action at a site, as well as convey information on the impact of taphonomic disturbance.

As mentioned earlier, human activities generally tend to produce a majority of splinters, whereas assemblages deposited by carnivores are frequently characterized by a high

91 proportion of long-bone cylinders with deleted epiphyses (Binford 1981:171–177; Bunn

1983a, 1989; Villa et al. 2004).

Several measures of fragmentation were used in this analysis. First, maximum length and maximum width were measured for all identified specimens. Length of all unidentified specimens was also recorded using 1 cm size classes. The circumference of long bone fragments was described as follows: less than half of the original circumference (<1/2), more than half (>1/2), or complete (Bunn 1983a; Villa and Mahieu

1991). Similarly, all long bone shafts were coded as a fraction of the original length: less than one quarter (<1/4), between one quarter and one half (1/4–1/2), less than three quarters (1/2–3/4), and three quarters to complete (>3/4) (Villa and Mahieu 1991).

Finally, long bone fragments were recorded using a simple system of five bone portions: proximal end, proximal shaft, middle shaft, distal shaft, and distal end (Marean and

Spencer 1991).

5.5.5 Burning

Burned bones are generally considered direct evidence of anthropogenic activity

(Stiner et al. 1995; Villa et al. 2004). Intentional burning of bones can result from cooking, disposal of rubbish in fires, use of bones as fuel, or cremation (Gilchrist and

Mytum 1986; Nicholson 1993; Villa et al. 2004). Bones can also be burned accidentally in a natural fire or due to their proximity to a fireplace (Gilchrist and Mytum 1986; Villa et al. 2004).

Burned bones can exhibit different colors, these being a function of the maximum temperature reached by the bone (Nicholson 1993; Lyman 1994:385; Stiner et al. 1995).

Bones that are slightly burned tend to be brownish. When reaching carbonization, they

92 turn black. As the intensity of the fire increases to calcination, bones shift from black to gray to white, sometimes passing through shades of blue or green (Shipman et al. 1984;

Gilchrist and Mytum 1986; Nicholson 1993; Stiner et al. 1995).

Burned bones tend to be more brittle than unburned bones. Their mechanical strength varies according to the extent to which they have been burned. Indeed, as burning intensity increases, bone become more friable and porous (Gilchrist and Mytum

1986; Stiner et al. 1995). Consequently, fragment size can be an important variable in determining burning intensity (Stiner et al. 1995). As bones become more susceptible to fragmentation, they may lose some of their diagnostic features, hindering taxonomic and skeletal identification. Bones will not necessarily be destroyed by burning, but rather may become analytically absent as they slowly become unidentifiable (Lyman and O'Brien

1987; Buikstra and Swegle 1989; Lyman 1994:391; Stiner et al. 1995).

In this study, the color of burned bones (brown, black, gray, green, blue, or white) was recorded for the entire assemblage. This method was chosen because colors are easily discernible to the naked eye and have proven reliable for diagnosing burning damage on archaeological bones (Stiner et al. 1995). The maximal length in centimeters of all burned fragments was also recorded to document fragmentation of the burned portion of the assemblage.

5.5.6 Additional taphonomic agents

Post-depositional processes may have an impact on the preservation of faunal assemblages and affect the identification of marks left on bones. Several types of alterations were recorded for all specimens: exfoliation, sheeting, root etching, staining, and cracking. Exfoliation can be defined as the loss of the first few millimeters of cortical

93 bone, while sheeting describes the fracturing of cortical bone into one or more sheets that tend to be parallel to the cortical surface (Morin 2012b:70). The six weathering stages identified by Behrensmeyer (1978) were not used in this study, because the criteria of cracking, flaking, and splintering used to categorize bone preservation were seldom observed on the Pacbitun assemblages. This is probably explained by the significant differences existing between the Belizean tropical rainforest and the Kenyan savannah where Behrensmeyer’s experiments were conducted.

In a different study, Tappen (1994) suggested that bone modifications caused by weathering in a savannah climate differ from those in a tropical rainforest. She observed that subaerial weathering, which caused the most important damage to the bones studied by Behrensmeyer, was slower in rainforests. She attributed this to the reduced sunlight, high moisture, and low variation of humidity and temperature distinctive of tropical rainforests. She occasionally observed exfoliation on the cortical surface of the bones, which is generally attributed to the chemical action of natural acidic soils and biological agents, such as roots, algae, and fungi (Tappen 1994; Fernández-Jalvo et al. 2002). This observation applies particularly to the Pacbitun assemblages, as it is the most common damage observed on the faunal remains.

As Tappen does not provide a methodology to study specifically bones recovered in tropical environments, bone surface condition was observed and assigned to one of four categories: intact, slightly damaged, damaged, or poor (Morin 2012b:71). In this classification, a surface is considered intact when almost no surface damage is recorded.

A surface with superficial damage is considered slightly damaged. This would be the case for a bone surface with local damage, but with visible marks or morphological features.

When landmarks are faint and the surface is significantly altered, the specimen is coded

94 as damaged. Poorly preserved bones have a considerably damaged cortical surface. Marks on this type of surface are unlikely to have survived. Finally, an estimate of the percentage of the remaining original bone surface was also recorded by 10% intervals

(Morin, personal communication, 2012).

5.6 Summary

The methods described in this chapter, which are used in the analysis of the

Pacbitun assemblages, will provide detailed taphonomic and quantitative data about the foraging strategies adopted by the Preclassic Maya at Pacbitun. The next chapter examines the stratigraphic and taphonomic integrity of the study sample.

CHAPTER 6: SAMPLE DESCRIPTION AND TAPHONOMY

This chapter assesses the integrity of the Pacbitun faunal assemblages. It begins with a description of the material analyzed in this study, while the second half focuses on measuring the impact of different taphonomic processes on the Middle Preclassic samples.

6.1 The Pacbitun faunal assemblages

The faunal assemblages examined in this study date exclusively to the Middle

Preclassic period (900–300 BC). The animal remains that were analyzed were recovered in Plazas A, B, C, and D during the excavations conducted from 1995 to 1997 by the

Trent University-Preclassic Maya Project and from 2008 to 2011 by the Pacbitun

Regional Archaeological Project (PRAP). The material recovered during the 1995 and

1996 field seasons has previously been identified by Norbert Stanchly. However, it was decided that this material would be reanalyzed in this study because different methods were used to assess the impact of taphonomic processes. The samples were retrieved from both primary (floor and perimeter deposits) and secondary contexts (fill and midden)

(Table 6.1).

The assemblages of vertebrate remains recovered from Pacbitun include a total of

1,730 specimens, with 566 specimens dating from the early Middle Preclassic and 1,164 from the late Middle Preclassic (Table 6.1). Most of the assemblages consist of unidentifiable mammal fragments (n = 1,110) and indeterminate specimens (n = 328).

The use of refitting during analysis reduced the initial NISP count by 39.0%, for a post- refit total of 292 identified specimens (Table 6.2). Only post-refit NISP counts are used in

96 the following analysis because they help to mitigate problems of differential fragmentation and identification, as well as minimize the problem of specimen interdependence (Morin 2012b:75). As such, post-refit NISP constitutes a more accurate estimate of taxonomic composition and skeletal part representation than pre-refit NISP.

Table 6.1 Number of specimens by primary and secondary contexts for the Middle Preclassic assemblages.

early Middle Preclassic late Middle Preclassic Total Contexts n n n Primary contexts Floor deposits 451 357 808 Perimeter deposits 84 139 223 Secondary contexts Plaza fill 2 26 28 Construction fill 29 21 50 Secondary midden 0 621 621 Total 566 1164 1730

Table 6.2 Pre- and post-refit NISP counts by time period.

Time period Total bone Pre-refit NISP* Post-refit NISP count n n early Middle Preclassic 566 210 125 late Middle Preclassic 1164 269 167 Total 1730 479 292

* Pre-refit counts include all refitted fragments.

Mammals are the most frequently identified animals in the samples (96.8% of the total assemblages, Tables 6.3 and 6.4). Artiodactyls, including the white-tailed deer, peccary, and red brocket deer, dominate both the early Middle Preclassic (56.8%) and late

Middle Preclassic samples (66.5%). Armadillos are the second most abundant taxon

(12.0%). Carnivores, represented by the domestic dog, coati, weasel, and various felines

(jaguar, puma, margay, and ocelot), represent 4.0% and 3.6% of the early Middle

Preclassic and late Middle Preclassic samples, respectively. Other identified mammals collectively form less than 8.0% of the assemblages and include the pocket gopher, opossum, tapir, paca, agouti, and rabbit. It should be noted that the collared peccary

(Pecari tajacu) and white-lipped peccary (Tayassu pecari) are considered difficult to differentiate osteologically (Olsen 1982; Emery 2010:64). The limited comparative collections available to the author did not permit taxonomic distinction between the two species.

Table 6.3 Distribution of faunal remains by zoological class for the early and late Middle Preclassic samples at Pacbitun.

Taxon early Middle Preclassic late Middle Preclassic Total n % n % n % Mammals 405 94.8 952 97.6 1357 96.8 Reptiles 14 3.3 17 1.7 31 2.2 Fish 6 1.4 2 0.2 8 0.6 Birds 2 0.5 3 0.3 5 0.4 Amphibians 0 0.0 1 0.1 1 0.1 Total 566 100.0 1164 99.9 1730 100.0

Reptiles form the second most important class (2.2%) in the assemblages. This group is dominated by turtle (early Middle Preclassic = 8.8%; late Middle Preclassic =

3.6%). Most turtle remains are carapace or plastron fragments lacking taxonomically diagnostic features. Only one specimen was specifically identified as a mud turtle. Snakes

(3.1%) represent the second most common reptile group, with six colubrid and three viper remains. Remains of iguanas (1.7%) constitute the only specimens of lizards identified in the samples.

Table 6.4 Identified taxa by NISP and MNI for the early and late Middle Preclassic samples at Pacbitun.

early Middle Preclassic late Middle Preclassic Scientific Name Common Name NISP % MNI % NISP % MNI % Osteichthyes Ictaluridae Catfish 1 0.8 1 4.2 - - - - Serranidae Grouper 1 0.8 1 4.2 - - - - Sparisoma spp. Parrotfish - - - - 1 0.6 1 4.2 Unidentified fish 4 3.3 - - 1 0.6 - - Amphibia Rhinella marina Marine toad - - - - 1 0.6 1 4.2 Reptilia Iguanidae Iguana 2 1.6 1 4.2 3 1.8 1 4.2 Testudines Turtle 11 8.8 1 4.2 5 3.0 - - Kinosternon spp. Mud turtle - - - - 1 0.6 1 4.2 Colubridae Colubrid 1 0.8 1 4.2 5 3.0 1 4.2 Viperidae Viper - - - - 3 1.8 1 4.2 Aves Galliformes Turkey, guan - - - - 1 0.6 1 4.2 Unidentified bird 2 1.6 1 4.2 2 1.2 - - Mammalia Didelphis marsupialis Common opossum - - - - 1 0.6 1 4.2 Didelphis virginiana Virginia opossum 2 1.6 1 4.2 1 0.6 1 4.2 Nine-banded Dasypus novemcinctus 17 13.6 4 16.7 18 10.8 2 8.3 armadillo Canidae Dog, fox 1 0.8 1 4.2 - - - - Canis lupus familiaris Domestic dog - - - - 2 1.2 1 4.2 Nasua narica White-nosed coati 1 0.8 - - - - Mustela frenata Long-tailed weasel - - - - 1 0.6 1 4.2 Felidae Cats 1 0.8 - - 1 0.6 - - Puma concolor Cougar, puma - - - - 1 0.6 1 4.2 Panthera onca Jaguar 1 0.8 1 4.2 - - - - Leopardus wiedii Margay - - - - 1 0.6 1 4.2 Leopardus pardalis Ocelot 1 0.8 1 4.2 - - - - Tapirus bairdii Tapir - - - - 1 0.6 1 4.2 Artiodactyla Artiodactyl 2 1.6 ------Tayassuidae Peccary 6 4.8 1 4.2 7 4.2 1 4.2 Cervidae Cervid 1 0.8 - - 6 3.6 - - Odocoileus White-tailed deer 51 40.8 2 8.3 88 52.7 3 14.8 virginianus Mazama americana Red brocket deer 7 5.6 1 4.2 10 6.0 1 4.2 Orthogeomys spp. Pocket gopher 4 3.2 2 8.3 3 1.2 1 4.2 Dasyproctidae Agoutis, pacas 1 0.8 ------Cuniculus paca Paca, gibnut 3 2.4 1 4.2 - - - - Dasyprocta punctata Agouti 2 1.6 1 4.2 1 0.6 1 4.2 Sylvilagus spp. Rabbit 2 1.6 1 4.2 2 1.2 1 4.2 Total 125 100.1 24 100.0 167 100.0 24 100.0

Fish remains were rarely encountered during analysis (0.6% of the total assemblages). The only identified specimen of freshwater species is a catfish vertebra, whereas marine species are represented by remains of grouper and parrotfish.

Identification of archaeological fish remains from Maya sites is generally considered challenging due to high species diversity, the non-diagnostic nature of many post-cranial fragments, and the propensity of fish remains to preserve poorly (Hamblin 1984; Powis et al. 1999; Stanchly 2004; Wake 2004a). Bird remains are also uncommon at the site (0.4% of the total assemblages). One specimen is identified as a gallinaceous bird. Although the high level of fragmentation precluded identification, the comparative material suggests the presence of medium- to large-sized birds. The amphibian assemblage is composed of a single bone from a marine toad.

Overall, mammals dominate the Middle Preclassic assemblages, while reptiles, fish, and birds only form a fraction of the material. Before discussing the significance of these observations regarding taxonomic composition, the second half of this chapter considers taphonomic processes that possibly affected the Pacbitun faunal samples.

6.2 Taphonomy

Taphonomy is a field of study that has received little attention from Maya archaeozoologists. In fact, published papers dealing specifically with the taphonomy of

Maya faunal assemblages are nearly nonexistent (for an exception, see Stanchly 2004) and taphonomy is rarely addressed in the studies of animal resource exploitation. The impact of taphonomic processes in assemblage formation is examined in dissertations by

Pohl (1976:67–81), Shaw (1991:243–249), and Emery (1997:72–92), but the scope of these analyses remains limited.

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This bias may be attributed to the generally accepted idea that unfavorable tropical environments are mainly responsible for poor preservation of organic remains at Maya sites. Although high soil acidity and humidity likely constitute the main factors responsible for the rapid deterioration of organic remains in the tropics, many other natural and cultural processes may also have impacted the preservation of faunal remains.

Therefore, as pointed out by Stanchly (2004), it is necessary to conduct detailed taphonomic analyses in order to identify the processes that might have shaped past faunal assemblages. The following section considers taphonomic factors relevant to the interpretation of the faunal samples at Pacbitun, including recovery methods, post- depositional attrition, burning, differential fragmentation and preservation, and carnivore ravaging.

6.2.1 Testing the stratigraphic sequence

Bone refits were used in this analysis to test the stratigraphic integrity of the faunal assemblages. A total of 81 refit sets were found in the Middle Preclassic assemblages, for an average of 3.3 specimens per set. Seventy-two of these sets are refits on dry-bone fractures and likely represent post-depositional breakage. Several specimens were also refitted on recent breaks probably produced during or after the excavations. All dry-bone refits are intra-level sets from the same unit. No anatomical refit was identified.

Because fractures on green bone are produced during the occupation of the site, they are more informative about site formation processes than dry-bone fractures (Morin et al. 2005). Nine sets of fragments involve green-bone fractures. Eight of them match specimens found within the same level of the same unit. Only one refit set on green-bone fracture is indicative of level disturbance, as it involves two fragments of a white-tailed

101 deer metacarpal recovered from an early Middle Preclassic deposit (Unit 2, Level 6b) and a late Middle Preclassic deposit (Unit 3, Level 4).

Overall, all but one refit involve fragments found in close proximity to one another. This suggests that post-depositional disturbance was limited at Pacbitun. This interpretation seems congruent with what was observed during the excavations at Plaza B.

Although the Maya had the habit of constructing new structures on top of older ones or modifying existing structures for new purposes (Garber et al. 2004a:25), the sub- structures at Plaza B do not seem to have been affected by this practice (T. G. Powis, personal communication, 2013). Indeed, the sub-structures appear to have been protected from later modifications by the thick midden that was laid over them at the end of the late

Middle Preclassic period. The presence of intact floor and alleyway surfaces (i.e., primary deposits) for both the early and late Middle Preclassic structures provides additional evidence for limited post-depositional disturbance. Additionally, no re-use of stones from the Middle Preclassic platforms for later constructions was observed (T. G. Powis, personal communication, 2013).

Concerning the late Middle Preclassic faunal remains recovered from the midden deposit, the small size of the artifact sample from this layer suggests a secondary nature

(Hohmann and Powis 1999; T. G. Powis, personal communication, 2013). Nevertheless, the covering of the plaza with construction fill during the Late Preclassic and Late Classic

(Hohmann and Powis 1996) seems to have protected the midden deposit from disturbance during later time periods.

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6.2.2 Recovery methods

The techniques used in the recovery of archaeological artifacts can affect sample composition in terms of species and skeletal representation, particularly when different screening methods are used (Payne 1975; Shaffer 1992; Emery 2004b). At Pacbitun, all deposits were dry-sieved in the field using a 1/4 inch (6.35 mm) mesh screen. However, excavation methods were not constant given that a 1/16 inch (1.2 mm) mesh screen was used to wet-sieve all floor deposits from Sub-Structures B-1 and B-2 in 2008 and 2009.

This variation in mesh size may affect the analysis of the Pacbitun samples. Indeed, studies have shown that the exclusive use of large sieves (i.e., 1/4 inch) typically creates a bias towards the recovery of large specimens (Casteel 1972; Shaffer 1992; Wake 2004b), whereas the use of finer mesh sizes often improves the recovery of smaller specimens, including that of fish, birds, and rodents (Shaffer and Sanchez 1994; Masson 2004b;

Wake 2004b; Serjeantson 2009). Therefore, it is necessary to consider the impact of the recovery methods on the abundance of small fragments at Pacbitun.

The difference between the samples sieved with 1/4 (6 mm) and 1/16 (1.2 mm) inch mesh screen is not statistically significant (Kolmogorov-Smirnov D = 0.33, p = 0.81,

Figure 6.1). This suggests that, although different recovery methods were used during the excavations, they did not influence the representation of small bone fragments. The size distribution of the fragments in Figure 6.1 also indicates that the Pacbitun assemblages are highly fragmented, with 74.2 % of the material classified as smaller than 2 cm, whereas less than 4.5% of the specimens are larger than 4 cm. The lowest size category

(<1 cm) may be artificially depressed, because faunal specimens of this size are often not systematically collected. This partly results from the difficulty of sorting bones smaller

103 than 0.5 cm, particularly when large quantities of minute bone fragments are present in faunal assemblages (Villa et al. 2004).

60 1/4 inch (n = 1278) 50 1/16 inch (n = 452) 40

20 % of specimens of % 10

0 <1 1–2 2–3 3–4 4–5 5+ Length in cm Figure 6.1 Fragment size distribution by screen size for all faunal specimens in the Pacbitun Middle Preclassic assemblages.

Frequencies of identified species for both sieve sizes were also compared (Table

6.5). Intuitively, one would expect to recover a higher frequency of small animals when a smaller screen mesh is used. However, at Pacbitun, the use of a 1/16 inch mesh screen did not lead to the recovery of a higher frequency of small animals, such as fish, birds, or rodents (D = 0.22, p = 0.95).

These results suggest that the use of a finer mesh size at Pacbitun did not result in an increased recovery of smaller fragments or taxa. However, other factors might have prevented the preservation of small specimens or species. These possibilities are explored in section 6.2.8 which provides a more detailed discussion of the taphonomy of fish and birds in the Pacbitun assemblages. In essence, it can be concluded that the use of two

104 different sampling strategies at Pacbitun has not greatly affected the assemblages in terms of fragment size and taxonomic composition.

Table 6.5 Taxonomic representation in the Middle Preclassic samples by mesh size, in percentages.

Taxon 1/4 inch (6 mm) 1/16 inch (1.2 mm) White-tailed deer 48.4 64.9 Brocket deer 6.9 5.4 Peccary 5.9 2.7 Armadillo 15.4 8.1 Agouti/paca 1.6 5.4 Reptiles 13.3 8.1 Small rodents 3.2 1.4 Fish 3.7 1.4 Birds 1.6 2.7 Total 100.0 100.0

6.2.3 Density-mediated attrition

In general, bones with low structural density are more prone to destruction by taphonomic processes than bones of higher mineral densities (Lyman 1994). This is because bone density is correlated with bone porosity. As the porosity of a given bone increases, so does the surface area per volume. As a result, mechanical and chemical attrition should affect more intensely bones of high porosity (or low density) simply because there is a greater surface to work on (Lyman 1982, 1994).

This observation is particularly important for this study, as the effects of acidic soils, water leaching, and plant and tree roots commonly observed on Maya faunal assemblages may preferentially damage and destroy bones of low density (Stanchly

2004). In addition, because their pores are filled with grease (Brink 1997), the spongy low-density portions of bones are attractive to humans and carnivores. These parts can possibly be removed from faunal assemblages by carnivore ravaging and human

105 activities, such as grease extraction and the use of bone as fuel (Binford 1981; Lyman

1982, 1994; Brink 1997; Morin 2010). Bone parts of high structural density are more likely to be targeted for object manufacture, because dense sections of bones are often preferred for tool and ornament production (Lyman 1982, 1994; Stanchly 2004; Emery

2008a, 2009).

In addition to anthropic and carnivore destruction, transport decisions may produce correlations with density, particularly when low-density elements (e.g., vertebrae) are discarded at kill sites (Binford 1978, 1981). For instance, vertebrae and crania were rarely encountered in the Pacbitun assemblages, but this might result from transport selectivity. Skulls may also have been preferentially used in the making of headdresses (Pohl 1981; Brown and Sheets 2000). To avoid this problem, only long bone portions were considered for examining density-mediated attrition in this study. Long bones are good candidates for this type of analysis because they are characterized by heterogeneous density, shafts usually being significantly denser than epiphyses.

Additionally, long bones should not be affected by transport decisions to the same extent as low-density elements, given that it is unlikely that long bone shafts and epiphyses were transported separately (Binford 1981).

To determine if density-mediated taphonomic processes have affected the

Pacbitun assemblages, frequencies of long bone portions of white-tailed deer were compared to density values. Although density values are available for deer (Odocoileus spp.), the method used by Lyman (1982, 1994) does not exclude the volume of the internal cavity and, therefore, significantly underestimates true bone density (Lam et al.

1999; Lam et al. 2003). Consequently, values for reindeer (Rangifer tarandus) were used

106 instead (Lam et al. 1999) (Table 6.6). Data from the early and late Middle Preclassic periods were considered together because of small sample size.

Table 6.6 Bone density values of Rangifer tarandus (Lam et al. 1999) compared to %NNISP and %MAU values for white-tailed deer long bone portions in the Middle Preclassic Pacbitun assemblages.

Bone portion Scan site Density (g/cm3) %NNISP %MAU Humerus proximal HU1 0.26 0.0 0.0 shaft HU3 1.12 100.0 100.0 distal HU5 0.48 37.5 50.0 Radius proximal RA1 0.53 25.0 33.3 shaft RA3 1.09 37.5 33.3 distal RA5 0.49 0.0 0.0 Ulna proximal UL1 0.49 37.5 33.3 shaft UL2 0.84 37.5 50.0 Metacarpal proximal MC1 0.92 75.0 83.3 shaft MC3 1.10 81.3 100.0 distal MC6 0.68 18.8 16.7 Femur proximal FE1 0.52 12.5 16.7 shaft FE4 1.15 62.5 66.7 distal FE6 0.32 62.5 83.3 Tibia proximal TI1 0.35 25.0 33.3 shaft TI3 1.13 50.0 50.0 distal TI5 0.73 12.5 16.7 Metatarsal proximal MR1 0.90 37.5 33.3 shaft MR3 1.08 43.8 50.0 distal MR6 0.59 18.8 16.7

Using Spearman’s rank order correlation coefficient (rs), both %NNISP and

%MAU values are positively and significantly correlated with reindeer bone density values (%NNISP rs = 0.61, p = 0.004; %MAU rs = 0.52, p = 0.02) (Figures 6.2 and 6.3).

These results suggest that the densest bone portions (shafts) of white-tailed deer were recovered more frequently than the less dense bone portions (epiphyses) from the same bones. Therefore, a destructive agent or process has probably affected the recovery of deer skeletal elements in the Pacbitun assemblages. It should be noted, however, that

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rs = 0.61, p = 0.004 100 sHUM

80 sMTC

pMTC sFEM dFEM 60

sTIB %NNISP sMT 40 pULN dHUM sULN pMTT sRAD

pTIB pRAD 20 dMT dMC pFEM dTIB

pHUM dRAD 0 0.00 0.20 0.40 0.60 0.80 1.00 1.20 Density (g/cm3) Figure 6.2 %NNISP of long bone portions of white-tailed deer versus bone density values (g/cm3). Data from Table 6.6. Abbreviations: p = proximal; s = shaft; and d = distal.

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rs = 0.52, p = 0.02 100 sMC sHUM

dFEM pMC 80

sFEM

60 sULN sTIB

%MAU dHUM sMT 40 pULN pTIB pRAD pMT sRAD

20 dMC pFEM dTIB dMT pHUM dRAD 0 0.00 0.20 0.40 0.60 0.80 1.00 1.20

Density (g/cm3)

Figure 6.3 %MAU of long bone portions of white-tailed deer versus bone density values (g/cm3). Data from Table 6.6. Abbreviations: p = proximal; s = shaft; and d = distal.

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19.8% of the indeterminate portion of the assemblages is made of spongy bone.

Therefore, cancellous bone has not disappeared from the assemblages. Instead, it is proposed that skeletal elements and element portions of low density have been fragmented by destructive processes to the extent that spongy specimens have become analytically absent from the samples (Lyman and O'Brien 1987; Morin 2010).

Differential preservation caused by bone density may not only affect skeletal part representation, but also the taxonomic composition of faunal assemblages. Unfortunately, it was not possible to perform analyses of bone density on smaller taxa due to small sample size and/or lack of bone density values.

In sum, the analysis of density-mediated attrition at Pacbitun simply indicates a correlation between bone density and element frequency. In order to identify the attritional agent(s) responsible for the destruction of spongy bones, the subsequent sections discuss bone burning, post-depositional destruction, and carnivore ravaging.

6.2.4 Bone burning

Because bones tend to become more friable as burning intensity increases (Stiner et al. 1995), it is necessary to assess the impact of burning on the Pacbitun material. At

Pacbitun, only a fraction of the assemblages (2.2%, n = 38) is affected by burning.

Burned bones are small, with no specimen larger than 4 cm. In fact, the majority (65.8%) are smaller than 2 cm. Although fragmentation might have affected bone identification, given that most burned bones are unidentified specimens, no difference in level of fragmentation was found between the burned and unburned portions of the assemblages

(D = 0.17, p = 0.999).

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Interestingly, none of the burned remains in the Pacbitun assemblages are from spongy bone. Many authors (e.g., Villa et al. 2004; Morin 2010) have argued that the preponderance of burned spongy bone in a faunal assemblage may be indicative of the use of bone as fuel. In contrast, cortical and spongy bone should be recovered in similar proportions if burning was used as a way to dispose of food waste (Clark and Ligouis

2010). Given the absence of burned spongy bone at Pacbitun, it seems unlikely that bone was used as fuel at the site. Overall, the low frequency of burned specimens seems inconsistent with burning as a major taphonomic agent in the Middle Preclassic assemblages at Pacbitun.

6.2.5 Post-depositional destruction

In order to assess the impact of post-depositional destruction, Marean (1991) has proposed a completeness index which estimates the fragmentation of small, compact elements, including the carpals, tarsals (with the exception of the calcaneus), lateral malleolus, and sesamoids. These bones were chosen by Marean because they are rarely fragmented by humans or carnivores for the extraction of nutrients. In rare instances, they may be fragmented for bone grease production (Binford 1978:164–165). These bones may also be swallowed by carnivores, but this does not seem of great concern at Pacbitun given that no digested remains were identified in the assemblages. Villa and colleagues

(2004) recommended the inclusion of the third phalanx in the index, because it should be more sensitive to post-depositional breakage given the small marrow cavity of this element. Following Villa and colleagues (2004), compact elements were coded as complete (CO), almost complete (ACO), or fragmented (FR). Bones with traces of percussion, chopping, or gnawing/digestion were excluded from the sample because their

110 fragmentation may not result from post-depositional destruction (Marean 1991). Both white-tailed deer and red brocket deer specimens were considered in the analysis.

The results of the completeness index (Table 6.7) may suggest that post- depositional breakage has not significantly altered the Middle Preclassic assemblages.

Indeed, the majority of the compact bones from both the early (66.7%) and late Middle

Preclassic (75.0%) samples are complete or almost complete. Nevertheless, the percentages also indicate that post-depositional breakage is responsible for at least some bone fracturing at the site. It should be noted that the index is influenced by the degree of identification of the remains. More complete specimens are easier to identify than fragmented ones, which may inflate the index of completeness. As a result, additional methods are required to assess the impact of post-depositional destruction on faunal assemblages.

Table 6.7 Degree of post-depositional completeness by time period.

Time period Total compact bones NISP CO+ACO %CO+ACO early Middle Preclassic 15 10 66.7 late Middle Preclassic 20 15 75.0 Total 35 25 71.4

Villa and colleagues (2004) argue that a high proportion (~70%) of green-bone fractures on long bones, combined with high values for the completeness index, is indicative of minimal post-depositional destruction. At Pacbitun, all long bone fragments from identified and indeterminate medium and large mammals were considered for analysis. Recent breaks were not included for obvious reasons. In the Pacbitun Middle

Preclassic samples, green-bone fractures are more common than dry-bone fractures, forming 69.8% (n = 104) of the assemblages (Table 6.8). The differences in the

111 frequencies of green-bone and dry-bone fractures between the early and late Middle

Preclassic samples is not statistically significant (χ2 = 0.01, p = 0.91).

Table 6.8 NISP counts for green- and dry-bone fractures by time period.

Fracture type early Middle Preclassic late Middle Preclassic Total n % n % n % Green -bone 36 69.2 68 70.1 104 69.8 Dry-bone 16 30.8 29 29.9 45 30.2 Total 52 100.0 97 100.0 149 100.0

The high frequency of fractures on green bone, as well as the results of the completeness index, is indicative of limited post-depositional breakage. It also suggests that a majority of long bone specimens were likely fractured before the deposition of the assemblages. To determine which agent is responsible for bone fracturing at Pacbitun, anthropic and carnivore activity at the site are discussed in the remainder of this chapter.

6.2.6 Bone surface preservation

Marks left on bones by carnivores and humans are critical for inferring the role of these two agents on assemblage formation, but surface damage may alter the presence of marks on faunal specimens. Therefore, before discussing anthropic and carnivore attrition, it is necessary to assess the preservation of bone surfaces at Pacbitun.

The preservation of the faunal remains ranges from poor to moderate (Table 6.9).

A majority of specimens (69.6%) display a poorly preserved surface, with few bones

(0.9%) considered as intact. This pattern is not unusual at Maya sites given that organic remains in the humid tropics do not appear to preserve well (Stanchly 2004; Wake

2004a). For instance, Emery (1997:315) reported that bone surfaces in the Petexbatun assemblages were highly eroded and, in many cases, had entirely disappeared.

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Table 6.9 Overall surface state for the Middle Preclassic Pacbitun assemblages.

Preservation state early Middle Preclassic late Middle Preclassic Total n % n % n % Poor 302 53.4 902 77.5 1204 69.6 Damaged 158 27.9 168 14.4 326 18.8 Slightly damaged 98 17.3 87 7.5 185 10.7 Intact 8 1.4 7 0.6 15 0.9 Total 566 100.0 1164 100.0 1730 100.0

The percentage of undamaged surface area was also recorded for all faunal specimens (Table 6.10). Each category of observable surface is represented in the two samples, but a majority of specimens are in damaged or poor condition. Fifty-five percent of the assemblages are characterized by a preserved surface area lesser than 20% of the initial bone surface, whereas only 12.9% have retained more than 60% of the original surface. This poor to moderate surface preservation may be partially attributed to the effects of weathering, given that exfoliation and root etching were observed on 94.7% and

21.3% of the fragments, respectively. Other less common alterations include staining

(6.2%), cracking (3.8%), and sheeting (1.0%). Overall, comparison of bone surfaces for the two samples suggests that the late Middle Preclassic material was more severely damaged by taphonomic processes than the early Middle Preclassic material. This is confirmed by statistical tests, given that differences in frequencies between the early and late Middle Preclassic samples are highly significant for both the overall surface condition (χ2 = 106.0, p < 0.0001) and the percentage of observable surface (χ2 = 174.5, p

< 0.0001).

The significant damage caused to the bone surfaces limited the taxonomic identification of the material, particularly long bone shaft fragments, by destroying

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Table 6.10 Percentage of observable surface in the Middle Preclassic samples.

Observable surface early Middle Preclassic late Middle Preclassic Total % n % n % n % 0–10 22 3.9 175 15.0 197 11.4 10–20 87 15.4 411 35.3 498 28.8 20–30 90 15.9 166 14.3 256 14.8 30–40 109 19.3 135 11.6 244 14.1 40–50 85 15.0 115 9.9 200 11.6 50–60 59 10.4 53 4.6 112 6.5 60–70 53 9.4 47 4.0 100 5.8 70–80 24 4.2 22 1.9 46 2.7 80–90 19 3.4 20 1.7 39 2.3 90–100 18 3.2 20 1.2 38 2.2 Total 566 100.0 1164 100.0 1730 100.0

muscle attachments and landmarks. Similarly, the poor surface preservation possibly hindered the identification of marks. Because of their faint nature, cutmarks may have been partly or entirely obliterated on a significant portion of the assemblages. Cutmarks were identified on only eight specimens at Pacbitun and were observed almost exclusively on specimens displaying a slightly damaged or damaged surface (Table 6.11).

An increase in cutmark frequency from the slightly damaged to damaged categories may indicate that these two categories of objects have possibly not been affected by surface damage to the same extent as the other two groups. The presence of only one specimen in the poor category, given that nearly half of the sample (49.3%) presents a poorly preserved surface, suggests that the total frequency of cutmarks may be somewhat underestimated in the assemblages. Taking these observations into consideration, the final section of this chapter examines the role of human and carnivore agents in the formation of the Pacbitun assemblages.

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Table 6.11 Frequencies of cutmarks on identified specimens and indeterminate long bone shafts by overall surface state for the Middle Preclassic samples at Pacbitun.

Preservation state n ncut %cut Poor 213 1 0.5 Damaged 117 5 4.3 Slightly damaged 94 2 2.1 Intact 8 0 0.0 Total 432 8 1.9

6.2.7 Human and carnivore agents

A priori, the Pacbitun faunal assemblages have been produced primarily by human activity. Most of the material was recovered inside or within the periphery of domestic structures (see Table 6.1) and found in association with ceramics, lithic tools, and shell beads (Arendt et al. 1996; Hohmann and Powis 1996, 1999; Hohmann et al.

1999; Powis 2009, 2010, 2011). Faunal remains recovered in secondary contexts (i.e., secondary midden, plaza fill, and construction fill) were also found with important quantities of domestic refuse. However, given the presence of domesticated dogs at Maya sites during the Middle Preclassic period (Pohl 1990; Shaw 1991; Wing and Scudder

1991; Clutton-Brock and Hammond 1994; Masson 2004a), this species may have modified or contributed to the faunal assemblages. Other carnivores, such as felines, mustelids, and procyonids, also have the ability to alter and accumulate bone assemblages

(Andrews and Evans 1983; Seymour 1989; Martín and Borrero 1997; Álvarez et al.

2012).

The presence of cutmarks, percussion notches, and burning on bone specimens

(Table 6.12) indicate that humans were involved in the formation of the Pacbitun faunal assemblages. Very few cutmarks (n = 8) were observed on the faunal specimens, but, as previously mentioned, this is likely the result of the generally poor preservation of bone

115 surfaces. This pattern seems typical of Maya sites, given that cutmarks are frequently identified on less than 1% of the faunal assemblages (e.g., Hamblin 1984:184–185; Carr

1986:293; Pohl 1990:157; Shaw 1991:226; Emery 1997:315). The cutmarks identified at

Pacbitun were found on the following skeletal elements: white-tailed deer femur (n = 1), white-tailed deer calcaneus (n = 1), white-tailed deer naviculocuboid (n = 1), white-tailed deer rib (n = 1), jaguar phalanx (n = 1), turtle shell fragment (n = 1), and mammal long bone fragments (n = 2).

Table 6.12 Frequencies of anthropogenic marks observed in the Middle Preclassic Pacbitun assemblages.

Time period Cutmark Percussion notch Burning n* total1 % n* total1 % n total % early Middle Preclassic 7 178 3.9 0 178 0.0 11 566 1.9 late Middle Preclassic 1 254 0.4 4 254 1.6 27 1164 2.3 Total 8 432 1.9 4 432 0.9 38 1730 2.2

* n represents the number of specimens displaying at least one cutmark or percussion notch. 1 the total includes identified specimens and indeterminate long bone shafts only.

Burned bones, an unambiguous marker of anthropogenic activity, are rare at

Pacbitun, representing only 2.2% of the total assemblages. Similar percentages were observed at Copán (2.6%; Collins 2002), while other Maya sites present higher values ranging from 5–16% (e.g., Hamblin 1984:185; Carr 1986:304; Shaw 1991:227; Emery

1997:315). Most of the burned bones in the Pacbitun assemblages are of brown or black color suggesting fires of moderate temperatures (Nicholson 1993; Stiner et al. 1995).

White or blue bones, more likely indicative of higher fire temperatures (Nicholson 1993;

Stiner et al. 1995), were rarely encountered. This observation applies to both the identified and indeterminate portions of the assemblages.

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The presence of notches was recorded during the analysis. However, due to overlap in notch morphologies produced by humans and carnivores (Capaldo and

Blumenschine 1994; Galán et al. 2009), they were not strictly considered diagnostic of human action. None of the four observed percussion notches were associated with gnaw marks at Pacbitun, which might suggest that these were produced during marrow cracking. Combined with the above reservations, percussion notches are presented along with cutmarks and burning as possible evidence of human involvement in assemblage formation.

Carnivore remains are rare at Pacbitun, forming only 3.8% of the assemblages (n

= 11, see Table 6.4). No single species dominate this group composed of dogs, coatis, weasels, and various large and small felines. One specimen, a jaguar phalanx, shows cutmarks, perhaps indicative of skinning. This suggests that at least some carnivore specimens were accumulated by the Maya.

Gnawing marks (e.g., tooth pits, punctures, furrows, etc.) affect only 1.9% of the assemblages (Table 6.13). These results are similar to those observed at Colha and

Cozumel, where carnivore marks were identified on 1.2% (Shaw 1991:244) and 2.0%

(Hamblin 1984:186) of the total assemblage, respectively. However, the frequency of carnivore-modified bones increases to 6.5% when unidentified specimens are excluded.

The low incidence of carnivore marks in the indeterminate portion of the assemblages may be caused by increased fragmentation, most unidentified specimens (84.8%) being no longer than 2 cm. Surface preservation might also be responsible for this pattern, considering that the majority of small indeterminate specimens have a poorly preserved surface.

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Table 6.13 Frequencies of carnivores marks observed in the Middle Preclassic Pacbitun assemblages.

Time period Gnawing Digestion n total % n total % early Middle Preclassic 14 566 2.5 0 566 0.0 late Middle Preclassic 19 1164 1.6 0 1164 0.0 Total 33 1730 1.9 0 1730 0.0

No traces of digestion or rodent marks were identified. Given the high level of fragmentation and the poor to moderate preservation of the Pacbitun faunal assemblages, it is possible that the digested remains did not survive at the site. As a matter of fact, digested specimens are more vulnerable to taphonomic processes than intact specimens because of the corrosion caused by gastric acids (Holwitz 1990). Therefore, their frequency may be slightly underestimated at Pacbitun, although digested bones do not seem common at Maya sites (Shaw 1991:244).

A qualitative assessment of the extent of gnawing on bone surfaces suggests that carnivore ravaging was limited at the site (Table 6.14). Most carnivore marks were coded as marginal (75.8%), meaning that they were uncommon on the bone surface. They appear almost exclusively on spongy bones, such as the vertebrae and epiphyses of long bones. This pattern is expected given that carnivores are primarily attracted to greasy bone parts. It also suggests that carnivores may be responsible to some extent for the fragmentation of spongy bone observed at the site.

Overall, the data on cutmarks, percussion notches, and burning suggests that most of the remains at Pacbitun were brought to the site and processed by humans. The low incidence of gnawing marks and digested remains probably indicates that carnivores contributed minimally to the bone assemblages and did not significantly modify

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Table 6.14 Extent of carnivore gnawing on bone surfaces for the Middle Preclassic samples at Pacbitun.

Limited to one Time period Marginal Covered Total section n % n % n % n % early Middle Preclassic 11 78.6 2 14.3 1 7.1 14 100.0 late Middle Preclassic 14 73.7 2 10.5 3 15.8 19 100.0 Total 25 75.8 4 12.1 4 12.1 33 100.0

the material. In fact, it seems reasonable to argue that carnivores, domestic dogs in particular, had secondary access to faunal remains after these had first been exploited by humans. The combined presence of cut and gnaw marks on a white-tailed deer femur

(PAC-97BO-002) seems to support this idea (Figure 6.4). Several punctures and tooth pits were observed on the lateral condyle of the distal end of the femur, whereas a set of cutmarks were identified in the intercondylar fossa.

Figure 6.4 Gnaw and cut marks on the distal end of a right white-tailed deer femur. On the left: two cutmarks in the intercondylar fossa. On the right: view of the lateral condyle with gnawing marks.

0 1 cm 0 1 cm

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Prey size may also constitute an argument in favor of this hypothesis. Indeed, the majority of gnaw marks (28/33, 84.8%) were found on bones of large mammals, such as white-tailed deer, red brocket deer, and peccary. However, most carnivores present in the

Maya subarea, with the exception of large felines, do not prey on animals of this size.

Studies have even shown that dogs are rarely successful in catching and killing deer

(Marchinton 1994). Therefore, it is most likely that carnivores modified assemblages first created by human activity. Nevertheless, the possibility that carnivores may have contributed sporadically to the assemblages, particularly by introducing smaller taxa such as rabbits, agoutis, and opossums, cannot be ruled out.

6.2.8 Taphonomy of fish and birds

Up to this point, this chapter has mainly focused on the impact of taphonomic processes on mammalian remains, the most common taxonomic class identified at

Pacbitun. This is not surprising given that mammals form 96.8% of the Pacbitun assemblages. However, under comparable depositional circumstances, remains of birds and fish may not behave in a manner similar to mammal remains. As a result, this section discusses the taphonomic history of fish and bird specimens recovered in the Middle

Preclassic assemblages at Pacbitun.

Fish and bird bones are generally considered less robust than mammalian bones

(Lyman 1994; Higgins 1999). For instance, actualistic research has shown that bird and fish specimens have a tendency to weather more rapidly than mammal remains (e.g.,

Nicholson 1996; Behrensmeyer et al. 2003; Cruz 2008). Fish remains are sensitive to the action of gastric acids and acidic soils, particularly if they have been boiled or moderately burned (Wheeler and Jones 1989; Butler 1990; Lubinski 1996). Because of their thin

120 cortical surface, bird bones may be easily fragmented and even destroyed by carnivore ravaging (Dirrigl 2001; Cruz 2008; Serjeantson 2009), although they might appear less attractive to carnivores than mammal bones as a result of their low marrow content

(Serjeantson 2009). It is also considered difficult to identify evidence of cultural modifications on avian remains (Dirrigl 2001). Cutmarks are generally more difficult to recognize on small- and medium-sized birds, because cutting or chopping implements may not be needed to disarticulate small skeletons. Birds of small size may also not need to be butchered because they can be cooked whole. Such practice would not leave any marks on the bones (Serjeantson 2009).

Similar to mammalian bone, some elements of bony fishes tend to preserve better than others. Cranial elements are generally considered to weather faster than vertebrae

(Butler 1990; Butler and Chatters 1994; Lubinski 1996), with the exception of teeth and otoliths, which are some of the most durable elements of fish (Butler and Chatters 1994).

It is important to note that fish of different families may not be affected by taphonomic processes to the same degree as a result of differences in the size, shape, and density of their skeleton (Wheeler and Jones 1989; Butler and Chatters 1994; Lubinski 1996; Butler and Schroeder 1998). Concerning birds, bone density has been recorded using different methods (e.g., Higgins 1999; Dirrigl 2001; Broughton et al. 2007), but the ranking of skeletal elements from densest to least dense is inconsistent between different studies. In general, thin and flat bones, such as the sternum and skull, are considered more fragile than long bones (Serjeantson 2009).

Different criteria may be combined in order to distinguish between culturally and naturally deposited assemblages of fish (Butler 1993; Zohar et al. 2001) and birds

(Serjeantson 2009), including taxonomic diversity, specimen size, bone structural density,

121 skeletal element representation, spatial distribution of the remains, and presence of burning and cutmarks. Due to the small size of both fish (n = 8) and bird (n = 5) samples at Pacbitun, these methods could not be applied. The fish remains are mostly represented by centrum fragments which lack spines. This considerably limited identification. One bird remain is identified as the fibula of a Galliform, whereas other specimens consists of three long bone fragments and one sternum fragment. None of the fish or bird remains display traces of human or carnivore modification. All fractures on bird long bones were identified as dry-bone fractures or were classified as ambiguous.

Because of the small size of the fish and bird samples and the absence of carnivore or anthropic marks on them, it is difficult to be conclusive about their mode of introduction in the Pacbitun assemblages. It is possible that the low frequency of small birds and fish was caused by differential preservation of these remains over time. It is possible that fish and bird remains were affected by taphonomic processes to a greater extent. They also might have been discarded elsewhere at the site (Emery 2004a, 2008b).

For instance, the discovery of pits filled with fish at Colha led Shaw (1991:238) to suggest that fish may have been buried quickly and separately from other food refuse.

The presence of domestic dogs at Pacbitun may also have decreased the probability of recovering fish and bird specimens because these may have been deleted from the assemblages by carnivores. Finally, these small species may also be absent simply because they were rarely exploited by the ancient Maya.

6.3 Summary

This chapter has shown that the Middle Preclassic assemblages at Pacbitun have been shaped by several taphonomic processes. The use of different recovery methods

122 does not seem to have affected the taxonomic composition of the assemblages. In comparison, weathering has altered a majority of faunal remains. This situation has significantly impeded the identification of the material. It is suggested that fragmentation of spongy bone may have been caused by carnivore ravaging of human-accumulated debris, some post-depositional destruction, and perhaps burning. Humans were likely responsible for the accumulation of most remains in the Middle Preclassic samples, although carnivores possibly contributed and modified a small portion of the assemblages. Overall, the extent of bone surface damage seems to be the only taphonomic process that has affected the early and late Middle Preclassic samples differently. The next chapter examines the subsistence strategies associated with the exploitation of animal resources at Pacbitun.

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CHAPTER 7: RESULTS

This chapter examines how the ancient Maya of Pacbitun utilized animal resources during the Middle Preclassic period. The chapter first begins with a description of the assemblages in terms of taxonomic composition, skeletal part representation, mortality profiles, and seasonality. Using the central place foraging prey choice model presented in Chapter 4, this chapter investigates the foraging strategies adopted by the

Pacbitun Maya. Finally, the findings of this study are compared with the subsistence patterns observed at other Middle Preclassic sites.

7.1 Taxonomic composition

Species abundances in the Pacbitun assemblages are presented using NISP counts.

MNI counts were not used in this study because of small sample size. A highly significant correlation between NISP and MNE values (early Middle Preclassic rs = 0.95, p < 0.001; late Middle Preclassic rs = 0.90, p < 0.001) also suggests that the ranking of identified species provided by these two quantification methods are similar. Therefore, the use of both methods in the presentation of taxonomic composition would be redundant.

The Middle Preclassic assemblages at Pacbitun are quite diverse given their small size (less than 170 identified specimens per time period). The early Middle Preclassic assemblage is represented by 18 different taxa, while 21 taxa were identified in the late

Middle Preclassic sample (Table 6.4). The early Middle Preclassic assemblage is dominated by white-tailed deer (40.8%). Other ungulates, namely the red brocket deer and peccary, comprise 5.6% and 4.8% of the sample, respectively. In the early Middle

Preclassic sample, most specimens identified as large mammals (n = 39) are also presumed to be from white-tailed deer. Although many of these specimens are long bone

124 shafts and rib fragments lacking diagnostic features, they present characteristics, such as size and density, that are consistent with that of white-tailed deer elements.

The armadillo (13.6%) and turtles (8.8%) also form a significant portion of the assemblage. However, it should be noted that the use of NISP counts may overestimate the importance of these two taxa in the present case. Indeed, both species have a very large number of bones as a result of the shells that form their carapace (Emery 2007b).

These bony scutes tend to preserve well and are highly diagnostic (Soibelzon et al. 2012).

While carapace or plastron fragments constitute the majority of identified turtle specimens, the armadillo is mainly represented by long bone fragments. The armadillo also produced the second highest MNE (n = 13), which may suggest that it was an important small game for the Pacbitun Maya. Other small game include large rodents

(agouti and paca), which collectively represent 4.8% of the sample, and the pocket gopher

(3.2%). Rare species (less than 2% of the assemblage) include the Virginia opossum, rabbit, iguana, colubrid snake, and all carnivores (i.e., domestic dog, coatimundi, jaguar, and ocelot). Fish remains are also not abundant (6.9%). Two specimens were attributed to the Ictaluridae (freshwater catfish) and Serranidae (grouper) families. Similarly, bird remains are infrequent (1.6%) and could not be identified more precisely than the class level.

The late Middle Preclassic assemblage is similar in composition to the early

Middle Preclassic (Kolmogorow-Smirnov D = 0.21, p = 0.74). The white-tailed deer is the most common taxon, representing 52.7% of the sample. It is followed by the armadillo (10.8%), red brocket deer (6.0%), and peccary (4.2%). Like the early Middle

Preclassic sample, most remains identified as large mammals (n = 65) are probably from white-tailed deer. Turtles are much less common in this sample (3.6%). One specimen

125 was identified as a mud turtle, while most of the other specimens are heavily weathered, poorly diagnostic carapace fragments. The proportion of rodents is also lower in the late

Middle Preclassic assemblage, with agouti and pocket gopher representing only 0.6% and

1.8% of the sample, respectively. Other mammals are rare (<2%). Fish are also less common than in the early Middle Preclassic sample, with one specimen identified as a parrotfish and another only identified as indeterminate fish. In contrast, reptile remains are proportionately more abundant in the late Middle Preclassic assemblage (colubrid =

3.0%; viper = 1.8%; iguanas = 1.8%). Birds are again rare (1.8%), with one specimen identified as a gallinaceous bird. One marine toad was also encountered. Although the

Middle Preclassic assemblages are characterized by considerable taxonomic diversity, the following discussion focuses principally on white-tailed deer because of small sample size.

7.2 Skeletal part representation

This section describes skeletal part representation for the white-tailed deer and offers a general description of skeletal element frequencies for three other taxa: the armadillo, peccary, and brocket deer. Small sample size precluded a detailed discussion of the other identified species. Standardized NNISP counts (%NNISP) were obtained by dividing all identified elements by the highest NNISP count. %NNISP values are used because they facilitate comparisons of assemblages. MNE and MAU values are presented with NNISP counts, although they are not used in this study to investigate skeletal part representation. At Pacbitun, NNISP and MAU values show similar rankings, as is suggested by the highly significant correlation between NNISP and MAU values for

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white-tailed deer (rs = 0.96, p < 0.001). Therefore, the use of one method over the other

should result in similar interpretations.

Due to small sample size, the early and late Middle Preclassic assemblages were

combined in order to investigate skeletal patterns for white-tailed deer. The most common

elements are mandibular and maxillary teeth, which are present in similar proportions

(Table 7.1, Figure 7.1). It is not unexpected for teeth to be abundant in the samples even

though the Pacbitun assemblages are poorly preserved. Teeth are among the densest

skeletal elements (Reitz and Wing 2008:46) and can be easily distinguished from bone

Maxillary teeth Mandibular teeth Atlas Axis Cervical vertebrae Thoracic … Lumbar vertebrae Sacrum Rib Scapula Humerus Radius Ulna Carpals Metacarpal Innominates Femur Tibia Tarsals Metatarsal Phalanx 1 Phalanx 2 Phalanx 3 Large sesamoid 0 20 40 60 80 100

%NNISP Figure 7.1 White-tailed deer body part representation in the Middle Preclassic assemblages at Pacbitun. Data from Table 7.1.

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Table 7.1 Skeletal part frequencies for white-tailed deer in the Middle Preclassic assemblages from Pacbitun. Bone portions not shown have a frequency of zero.

Body portion NISP NNISP %NNISP MNE MAU %MAU Maxillary teeth 8 4.00 88.9 3 1.50 50.0 Mandibular teeth 9 4.50 100.0 4 2.00 66.7 Cervical vertebrae 2 0.29 6.3 2 0.29 9.5 Thoracic vertebrae 1 0.08 1.7 1 0.08 2.6 Lumbar vertebrae 1 0.17 3.7 1 0.17 5.6 Rib 7 0.27 6.0 5 0.19 6.4 Scapula 1 0.50 11.1 1 0.50 16.7 Humerus, shaft 8 4.00 88.9 6 3.00 100.0 Humerus, distal 3 1.50 33.3 3 1.50 50.0 Radius, proximal 2 1.00 22.2 2 1.00 33.3 Radius, shaft 3 1.50 33.3 2 1.00 33.3 Ulna, proximal 3 1.50 33.3 2 1.00 33.3 Ulna, shaft 3 1.50 33.3 3 1.50 50.0 Scaphoid 1 0.50 11.1 1 0.50 16.7 Lunatum 1 0.50 11.1 1 0.50 16.7 Triquetrum 2 1.00 22.2 2 1.00 33.3 Capitatum 1 0.50 11.1 1 0.50 16.7 Metacarpal, proximal 6 3.00 66.7 5 2.50 83.3 Metacarpal, shaft 5 2.50 55.6 5 2.50 83.3 Innominate 3 1.50 33.3 2 1.00 33.3 Femur, proximal 1 0.50 11.1 1 0.50 16.7 Femur, shaft 5 2.50 55.6 4 2.00 66.7 Femur, distal 5 2.50 55.6 5 2.50 83.3 Tibia, proximal 2 1.00 22.2 2 1.00 33.3 Tibia, shaft 4 2.00 44.4 3 1.50 50.0 Tibia, distal 1 0.50 11.1 1 0.50 16.7 Malleolar 3 1.50 33.3 3 1.50 50.0 Astragalus 5 2.50 55.6 4 2.00 66.7 Calcaneum 6 3.00 66.7 5 2.50 83.3 Naviculocuboid 6 3.00 66.7 5 2.50 83.3 Greater cuneiform 4 2.00 44.4 4 2.00 66.7 Metatarsal, proximal 3 1.50 33.3 2 1.00 33.3 Metatarsal, shaft 2 1.00 22.2 2 1.00 33.3 Metapodial, shaft 3 0.75 16.7 2 0.50 16.7 Metapodial, distal 3 0.75 16.7 2 0.50 16.7 Phalanx 1 9 1.13 25.0 6 0.75 25.0 Phalanx 2 8 1.00 22.2 7 0.88 29.2 Phalanx 3 3 0.38 8.3 3 0.38 12.5 Large sesamoid 5 0.31 6.9 5 0.31 10.4 Total 148 118

fragments even when heavily weathered. The rest of the sample is dominated by long bones, with shaft fragments being slightly more common than epiphyses. As discussed in

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Chapter 6, this situation probably results from mediated-density attrition. The humerus is the most common long bone, followed by the metacarpal and femur. Tarsals, and to a lesser extent carpals and phalanges, are also abundant. Cranial elements and mandibles are only represented by loose teeth. Elements of the axial skeleton, such as vertebrae and ribs, are rare in the assemblages. The under-representation of these elements may result from differential preservation, as elements of low structural density are less frequent in the assemblages than elements of higher density.

The peccary and red brocket deer have a respective NISP of 13 and 17 (Table 7.2).

As for the white-tailed deer, elements of the axial skeleton are poorly represented in these species. Long bone specimens are well represented for the brocket deer (7/17, 41.2%), but

Table 7.2 Skeletal part frequencies (NISP) of armadillo, peccary, and red brocket deer for the Middle Preclassic period at Pacbitun. Bone portions that are not listed have a frequency of zero.

Body part Armadillo Peccary Brocket deer Maxillary teeth 1 Cervical vertebrae 1 Scapula 1 Humerus 1 Radius 2 2 Ulna 1 1 Carpals 3 Metacarpal 1 2 Femur 2 1 Tibia 10 1 Fibula 2 Tarsals 2 2 2 Metatarsal 1 2 Phalanx 1 1 3 4 Phalanx 2 3 Phalanx 3 1 Scute 12 Total 35 13 17

129 not as abundant for the peccary (3/13, 23.1%). Phalanges, carpals, and tarsals are most common in these species (peccary = 8/13, 61.5%; brocket deer = 10/17, 58.8%). The armadillo is primarily represented by bony scutes (12/35, 34.2%). This pattern is not surprising given that these elements are highly diagnostic. Long bones are also frequent, particularly the tibia (10/35, 28.6%).

7.3 Mortality profiles

As it is the case for most faunal assemblages recovered from Maya sites, data concerning prey age and sex is very limited in the sample. Osteometric measurements could not be applied in this study due to the highly fragmented nature of the specimens. It should be noted that the usefulness of osteometric data in the tropics may be limited to few animals, including white-tailed deer, because many mammal species present in the study region, such as the peccary, rabbit, or agouti, do not exhibit any apparent or only show weak sexual dimorphism (Ojasti 1996).

Antlers may also provide coarse information on the sex and age composition of an assemblage. Unfortunately, antler remains are absent from the Pacbitun samples. This pattern may have several causes. For instance, the Preclassic Maya may have preferentially used crania (with attached antlers) for decorative and ritual purposes, for instance in the making of headdresses (Pohl 1981; Brown and Sheets 2000) or as offerings in hunting shrines (Brown 2005). Antlers may also have been left at kill sites because they are heavy to carry and possess little food value (Binford 1978). However, this observation may not apply to the present study. Because of the small size of white- tailed deer in the tropics (30–50 kg), small groups of foragers or even a single hunter may have been able to transport complete carcasses back to the site, including crania with

130 antlers. Lastly, a hunting strategy biased against male individuals could also have created this pattern.

Data on age profiles was gathered from the analysis of tooth eruption and wear and epiphyseal fusion for white-tailed deer. The only tooth dated to the early Middle

Preclassic was too fragmented for adequate measurement. In the late Middle Preclassic sample, two deciduous teeth were available for analysis. The first one is a lower fourth premolar. Crown height measurement, combined with the absence of dental wear, suggests an age of less than seven weeks (Severinghaus 1949). The second deciduous tooth is also a lower fourth premolar, which exhibits considerable wear. The age of this individual is estimated between 17–20 months, falling within the subadult category (1–2 years old) (Severinghaus 1949; Gee et al. 2002). The number of permanent teeth adequately preserved for crown height measurements is also relatively small. Three specimens are classified as subadults: two lower first molars correspond to an age at death of 20–26 months, whereas one second lower molar exhibiting little wear provides an age of about 13–14 months (Severinghaus 1949). Only one tooth, a lower third molar, is classified as pertaining to an adult (>2 years old). The presence of little to no wear (Gee et al. 2002) suggests the presence of a young individual. No completely worn teeth were identified in the Middle Preclassic assemblages, suggesting the absence of old and very old individuals.

Although it may only provide coarse information on mortality profiles, the age of white-tailed deer specimens was also determined using sequences of epiphyseal fusion

(Purdue 1983). The main drawback of this method is that it does not allow to extrapolate the age of an individual passed the time of bone fusion. For the early Middle Preclassic sample, 19 bones were analyzed (Table 7.3). From these, three specimens show unfused

131 epiphyses. The identification of these unfused bone portions indicate the presence of young individuals (<2 years old) in the assemblage. The presence of four fused first phalanges, which are completely fused between 17–20 months of age, indicates that subadults and adults may have been taken. Finally, ten out of nineteen specimens are element portions which are completely fused between 26–38 months of age. This suggests that about half of the specimens in the sample are from adult individuals.

Table 7.3 Number of specimens identified per category of epiphyseal fusion for white- tailed deer in the early and late Middle Preclassic samples at Pacbitun. Age of epiphyseal fusion is provided in months.

Unfused Intermediate Fused Time period Element Age1 n Age n Age1 n Radius, px 5–8 1 2nd phalanx 11–17 1 1st phalanx 17–20 4 Metacarpal 20–23 1 26–29 3 early Middle Preclassic Ulna, px 20 1 (19 specimens) Calcaneum 26–29 3

Femur, di 23–29 1 26–38 1 Tibia, px 26–38 1 Ulna, px 26–38 1 Cervical vertebra, ant 38 1 Radius, px 5–8 1 Coxal 8–11 1 2nd phalanx 11–17 7 Humerus, di 12–20 3 1st phalanx 17–20 5 late Middle Preclassic Tibia, di 20–23 1 (31 specimens) Metacarpal 20–23 1 26–29 2 Calcaneum 26–29 3 Metatarsal 26–29 3 Femur, di 26–38 3 Femur, px 32–38 1 1 Data on epiphyseal fusion in months from Purdue (1983). A range of values is provided when data for males and females differed. Abbreviations: px = proximal, di = distal, and ant = anterior.

Only one specimen in the late Middle Preclassic assemblage, a distal metapodial epiphysis which generally fuses at 20 months of age, is unfused (Table 7.3). Sixteen

132 specimens are from bones which fuse between 11–23 months of age, suggesting the presence of individuals which can be considered as subadults or full adults. Twelve additional specimens are completely fused between 26–38 months of age. Therefore,

40.0% of the late Middle Preclassic specimens would belong to the subadult or young adult category.

Published sequences of teeth eruption and wear are also available for the domestic dog, peccary, and paca. However, the information obtained for these taxa is very limited: no more than one or two teeth could be analyzed for each species. Based on the timing of dental eruption for collared peccary (Kirkpatrick and Sowls 1962), an upper third premolar from the early Middle Preclassic was identified as belonging to an individual at least 18 months old. However, the fact that it shows considerable wear suggests that the animal was much older. The only dog tooth identified in the samples is a lower first molar from the late Middle Preclassic period. This tooth erupts between three to five months of age (Silver 1969). Given the limited degree of wear observed on the cusps, the tooth is attributed to a juvenile individual. In the case of the paca, the presence of intact folds on a lower first molar, which become isolated early in the wear sequence (Ungar 2010:215), suggests a very young individual. Another paca tooth was identified as a lower third molar. Although this tooth erupts at about 14 months of age (Oliveira and Canola 2007), the heavy wear and isolated folds indicate a much older individual. These two teeth date to the early Middle Preclassic period.

Overall, data on tooth wear and epiphyseal fusion for white-tailed deer suggests that the Pacbitun Maya focused on the exploitation of subadults and young adults. Very old individuals appear to have been absent in the assemblages. However, this pattern may be an artefact of small sample size, given that only six teeth were available for analysis.

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There is also limited evidence for the exploitation of very young individuals. However, it should be noted that unfused epiphyses, as well as deciduous teeth, are more vulnerable to the action of post-depositional destruction or carnivore ravaging than completely formed bones or teeth (Klein and Cruz-Uribe 1984:43). Given that the Pacbitun assemblages have been affected to some degree by these two processes, very young individuals may be under-represented because of differential preservation.

7.4 Scheduling of activities

The Middle Preclassic residents of Pacbitun occupied a landscape which was spatially and seasonally diverse. Given that they largely subsisted on agriculture, the ancient Maya would have had to integrate animal resource procurement into the scheduling of agricultural activities throughout the year. Ethnographic accounts suggest that swidden agriculture, the type of agriculture which is believed to have been practiced at Pacbitun during the Middle Preclassic (Healy et al. 2004b), was more likely carried out during the wet season, although wetlands can also be cultivated during the dry season

(Pohl 1990:155). Given that most species in the tropics can breed year-round, or have birthing peaks extending over several months (Chapter 4), it was not possible to determine the season of capture of the species identified at Pacbitun. This situation inhibited insights regarding activity scheduling.

7.5 Diet breadth

The investigation of diet breadth can provide information about past foraging efficiency and animal procurement strategies. At Pacbitun, diet breadth was examined by comparing taxonomic abundances using the prey ranks presented in Chapter 4. According to the body size rule (Figure 4.1), the tapir should be considered the highest-ranked prey

134 type. At Pacbitun, this animal is only represented by a single specimen. Following the predictions of the central place forager prey choice model, it is suggested that the scarcity of tapir possibly reflects a decline in the abundance of this animal in patches located near the site. Consequently, the residents of Pacbitun should have invested more time in searching for and handling resources ranked lower on the body mass scale (see discussion about artiodactyls). It is also possible that this animal was infrequently encountered because of its behavioral habits. Indeed, tapirs are solitary animals, which travel extensively and prefer to live in the dense, low vegetation of mature forests. In fact, this animal is seldom encountered at Maya sites. Ethnohistoric and ethnographic accounts report that the tapir is not usually taken by the Maya; it is either regarded as too difficult to kill and transport (Tozzer 1941:203; Jorgenson 1999) or as non-edible (Hopkins 1992;

Fry 2009).

The jaguar and puma occupy the second and third ranks on the body mass scale

(Figure 4.1). Although predatory defense mechanisms make carnivores dangerous to exploit in comparison to the relatively docile white-tailed deer, tapir, or peccary, this parameter may explain why felines would have been taken by the Maya. Indeed, pursuing large, high-risk animals may not always be the most profitable activity in terms of return rates, but it can confer prestige onto hunters (McGuire and Hildebrandt 2005; Lupo

2007). At Pacbitun, felines are not abundant in the Middle Preclassic assemblages (early

Middle Preclassic = 2.4% of NISP, late Middle Preclassic = 1.8%). However, carnivores are generally encountered less frequently than herbivores as a result of their high rank in the trophic chain.

Evidence from the Classic period suggests that bones of large felines were used in the manufacture of objects (Moholy-Nagy 1994; Emery 2008a, 2009) and that their skins

135 and teeth were worn by priests and rulers as a symbol of their high social status (Pohl

1983; Hopkins 1992; Emery 2010). It is not known if the animals were consumed during the Classic period. Felines may have been used in a similar fashion during the Middle

Preclassic. Skeletal patterns may provide evidence for the exploitation of pelts given that phalanges, metapodials, and caudal vertebrae may be left with the pelt when fur-bearing animals are skinned (Fairnell 2008; Reitz and Wing 2008:127). At Pacbitun, four out of six feline elements fall within this category: two phalanges from a jaguar and a small felid, one navicular from a margay, and one caudal vertebra from a small felid. Two humeri from a puma and an ocelot complete the assemblage. Although this skeletal pattern seems to point towards the exploitation of pelts, the jaguar phalanx displays three sets of deep cutmarks which probably result from the disarticulation rather than the skinning of the foot. Overall, the very small sample size of the feline assemblage makes it difficult to reach conclusions about the exploitation of these animals at Pacbitun. It seems probable that humans were involved in the accumulation of felines during the Middle

Preclassic, although these taxa are only present in limited proportions at the site.

Four other ungulates are also considered high-ranked resources. These are the white-tailed deer, red brocket deer, white-lipped peccary, and collared peccary. Following

Emery (2008b), an abundance index was calculated to evaluate the importance of these large animals relative to smaller taxa. This “artiodactyl index” was calculated as follow: Ʃ artiodactyls / Ʃ (artiodactyls + all other mammals excluding tapir, jaguar and puma).

During the early Middle Preclassic, hunting strategies seem to have focused on the exploitation of the four ungulates, as suggested by a value of 0.65 for the artiodactyl index. Higher values were observed for the late Middle Preclassic sample (artiodactyl index = 0.77), but the difference in proportions of artiodactyls between the two time

136 periods is not statistically significant (K-S D = 0.33, p = 0.97). From these four taxa, the white-tailed deer was the species most commonly targeted by the Middle Preclassic

Maya. This taxon comprises 79.7% of the artiodactyl sample from the early Middle

Preclassic period and 83.8% of the late Middle Preclassic artiodactyl sample. It is suggested that the Pacbitun Maya concentrated their efforts on the procurement of these four artiodactyls because higher ranked prey types were infrequently encountered.

Many of the mammals of small and intermediate size, which are ranked low on the body mass scale, are rare in both the early and late Middle Preclassic samples. This pattern is, with the exception of the armadillo, in accordance with the prey ranking. It is possible that the high frequency of armadillo does not relate to the amount of meat it can procure, but rather to the amount of fat it can provide. During the wet season, armadillos build an extremely thick layer of fat relative to their body size (Hill et al. 1984; Hill et al.

1987). This characteristic could make them a desirable prey type in comparison to other small- and medium-sized mammals that tend to be much leaner. Unfortunately, the lack of quantitative data on body fat does not allow the testing of this hypothesis. Other taxa that likely violate the body size rule include turtles. These were exploited at Pacbitun, although their high frequency in the samples may be exaggerated by the use of NISP counts. Nonetheless, turtles were possibly always included in the optimal diet. As slow- moving animals, they provide important quantities of meat for little handling costs. Birds and fish seem to have been considered as resources of low profitability given that they are rare at the site.

In Chapter 4, it was discussed that the en masse collection of small animals may have created rank inversions. Given the low abundance of nearly all small prey items at

Pacbitun, the Middle Preclassic Maya do not appear to have invested much time in this

137 hunting strategy. Similarly, certain animals may have been preferentially targeted because their gregarious behavior increases the chances of handling multiple individuals during a foraging bout. Two species identified at Pacbitun live in large groups: the peccary and coati. Given that only one specimen of coati was identified in the samples, the gregariousness of this medium-sized animal does not appear to have modified its low ranking on the body mass scale. According to the prey ranking, the peccary is an animal of high utility at Pacbitun. Therefore, its gregarious behavior would not have affected its ranking given that it is already included in the optimal diet.

The presence of snakes at Pacbitun raises the question of whether they were exploited by the Maya. Snakes are known to have been used as offerings in sacrificial ceremonies or as ornaments by the Classic period Maya (Pohl 1990; Emery 2010). They might also have been used as a source of food. Unfortunately, none of the snake specimens identified at Pacbitun present traces of anthropic activity. The presence of snakes at archaeological sites is often considered to be incidental because they are attracted to human habitations and may die there naturally. However, given that snakes are not scavengers, they do not naturally dwell in middens or refuse piles. Fradkin (2004) suggests that their presence in important quantities in this type of deposits likely result from anthropic activity. This information is crucial for the interpretation of snake exploitation at Pacbitun given that a majority of snake specimens (66.7%) were recovered in the secondary midden dating to the late Middle Preclassic period. Additionally, the size of two viper vertebrae indicates the presence of large snakes which could have provided sizeable quantities of meat. In sum, although evidence for the exploitation of snakes at

Pacbitun is conjectural, it is suggested that the Middle Preclassic Maya may have intentionally captured and exploited these reptiles.

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The only amphibian remains recovered at Pacbitun is that of a marine toad. The presence of this species in the assemblage is also difficult to interpret. It is not clear if this giant toad was eaten by the Maya because its parotoid glands produce secretions which are toxic (Campbell 1998:68). Marine toads thrive in disturbed habitats and are often found near human settlements (Campbell 1998:69). As a result, the specimen identified at

Pacbitun may be intrusive or perhaps was used as poison or hallucinogenic drug.

At this point, the discussion of diet breadth has focused on hunted animals.

Domestic dogs are known to have been used by the Maya during the Middle Preclassic period. These animals likely served as pets and hunting companions, but evidence from sites in northern Belize suggests that they were also used as a source of food (Shaw 1991;

Wing and Scudder 1991; Clutton-Brock and Hammond 1994; Masson 2004a). This practice seems unlikely at Pacbitun given that only three specimens of Canidae were recovered at the site. However, the possibility that some dogs were given a mortuary treatment different from that of other taxa cannot be discounted (Losey et al. 2011). For instance, at Cuello, dogs were recovered in special deposits and interpreted as having been used in sacrifices (Tykot et al. 1996; White et al. 2001b). Therefore, it is possible that dog remains were not frequently recovered near residential structures or in middens at Pacbitun because their remains were buried elsewhere.

In summary, the taxonomic data suggests that diet breadth at Pacbitun during the

Middle Preclassic was, on average, relatively narrow. Procurement of animal resources focused on white-tailed deer. Other artiodactyls were exploited in much lower proportions. The highest ranked taxa, namely the tapir, jaguar, and puma, appear to have been infrequently encountered. The available data demonstrate that a variety of lower- ranked resources were also included in the diet, but their low abundances in the samples

139 suggest that they were only occasionally taken. Although the early and late Middle

Preclassic samples are small in size, they both offer a similar picture of animal resource procurement (K-S D = 0.21, p = 0.74).

7.6 Habitat use

The species identified in the Pacbitun vertebrate assemblages can inform us on the types of habitats exploited by the Middle Preclassic Maya. Generally, information derived from analyses of habitat use is coarse in nature because many tropical animals can thrive in a variety of habitats. The use of habitat fidelity statistics may help to overcome this limitation because values are assigned to animal species as a function of their fidelity to different types of habitats. Habitats considered in this analysis include closed canopy forest, secondary/disturbed forest, riverine/lacustrine habitats, agricultural/open areas, and residential areas (Table 7.4). In this study, the NISP count of identified species was multiplied by values of habitat fidelity obtained from Emery and Thornton (2008b). It should be noted that this analysis only includes mammalian taxa because of a lack of data and/or less precise taxonomic identification for birds, reptiles, and fish in the Pacbitun assemblages. Although these animals are excluded from the analysis of habitat fidelity, they are included in the general discussion of habitat use.

The analysis of habitat fidelity suggests that the Pacbitun Maya focused their efforts on the procurement of animals which mainly thrive in secondary forests (41.3%) or agricultural fields (36.8%, Table 7.5). These results are not unexpected. At Pacbitun, the practice of swidden agriculture during the Middle Preclassic period likely resulted in the partial deforestation of the landscape. This farming strategy generally leads to the formation of areas of second growth vegetation which act as a transition between

140 undisturbed canopy forests and open cultivated areas (Emery 2010). The majority of mammals identified in the assemblages might have been attracted to this patchy environment because they can find shelter in second-growth forests during the day and visit agricultural fields and household gardens at night to feed on crops, fruit trees, and other greens. These animals include the white-tailed deer, collared peccary, and smaller game, such as the agouti, opossum, armadillo, and rabbit.

Table 7.4 Habitat fidelity values for the mammalian species identified in the Middle Preclassic assemblages at Pacbitun. Values from Emery and Thornton (2008b:164).

Taxon MF SEC RIV AGR RES Opossum 0.1 0.5 0.2 0.2 Armadillo 0.2 0.4 0.4 Coatimundi 0.35 0.4 0.2 0.1 Long-tailed weasel 0.4 0.4 0.2 Cougar 0.4 0.4 0.2 Jaguar 0.65 0.2 0.2 Margay 0.6 0.4 Ocelot 0.5 0.3 0. 2 Tapir 0.2 0.8 Peccary 0.6 0.2 0.2 White-tailed deer 0.1 0.5 0.45 Red brocket deer 0.6 0.3 0.1 Pocket gopher 0.5 0.5 Paca 0.45 0.5 0.1 Agouti 0.25 0.3 0.4 0.1 Rabbit 0.5 0.5 Abbreviations: MF = mature/closed canopy forest, SEC = secondary/disturbed forest, RIV = riverine or lacustrine habitats, AGR = habitats with low arboreal vegetation, including savannas and agricultural fields, and RES = residential areas.

Table 7.5 Analysis of habitat fidelity for the Pacbitun assemblages, by time period. Results are provided in percentages.

Time period MF SEC RIV AGR RES Total early Middle Preclassic 20.5 40.9 2.4 35.7 0.5 100.0 late Middle Preclassic 18.3 41.6 1.9 37.9 0.3 100.0 Total 19.4 41.3 2.1 36.8 0.4 100.0

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Animals which occupy closed forested areas also seem to have been acquired

(19.4%), but not to the same extent as animals which thrive in disturbed habitats. This result is consistent with the low abundance of forest-dwelling animals in the assemblages, such as the white-lipped peccary, red brocket deer, paca, jaguar, margay, and tapir. These animals do not adapt well to human presence and are most commonly found in undisturbed dense forests. Therefore, the presence of these species in the Middle

Preclassic assemblages suggests that hunting trips were at least occasionally taken to forested habitats.

Riverine habitats do not appear to have been extensively exploited (2.1%). This is unexpected given that many secondary and tertiary water sources are available within 5–

10 km of Pacbitun. This result is supported by the low abundance of freshwater fish in the assemblages. However, it should be noted that the exclusion of reptiles from the analysis of habitat fidelity may distort the picture. Indeed, iguanas spend most of their time in trees found near permanent sources of water, whereas turtles can be found in a variety of freshwater habitats. The very large quantities of invertebrate remains recovered in the

1995 and 1996 excavations also provide evidence for the exploitation of freshwater habitats (Stanchly 1999). Unfortunately, the dietary importance of the identified species, namely the jute snails, apple snails, and pearly mussels, remains poorly understood

(Powis 2004; Solis 2011). In sum, the combined presence of fish, iguanas, and turtles in the assemblages suggests that the Pacbitun Maya exploited riverine resources, perhaps in proportions slightly higher than that suggested by the habitat fidelity analysis.

There is also evidence at Pacbitun for the procurement of resources from non-local habitats. The identification of marine fish, such as the parrotfish and grouper, is indicative

142 of long-distance trade or the exploitation of the Caribbean coast, located 150 km east from Pacbitun.

Overall, animal procurement strategies during the Middle Preclassic seem to have focused on the exploitation of local terrestrial habitats, with possible access to resources acquired from distant locales, such as the Caribbean Sea. Foraging strategies appear to have targeted prey types that could be acquired at little distance from the site, particularly in secondary growth forests or agricultural fields. Patterns of habitat use are similar for both the early and late Middle Preclassic periods (K-S D = 0.2, p = 0.999).

7.7 Transport selectivity

At Pacbitun, none of the species identified in the Middle Preclassic assemblages are represented by complete skeletons. As explained in Chapter 4, central place foragers who hunt large prey items often need to make decisions regarding which parts to bring home and which ones to leave behind. In order to investigate such decisions, archaeozoologists have developed indices of economic utility (e.g., Binford 1978;

Metcalfe and Jones 1988; Morin 2007; Morin and Ready 2013). These indices assume that different skeletal parts yield different amounts of meat, marrow, and grease. As a result, skeletal elements can be ranked according to their food value.

If foragers were highly selective about which parts to transport to a central place, skeletal elements of higher food utility should be more abundant in the archaeological record than those of low utility. However, if carcasses were brought whole, all skeletal elements should be present in roughly equal proportions. This observation may apply to three high-ranked species exploited at Pacbitun: white-tailed deer, red brocket deer, and peccary. Given that these large prey items all weigh less than 50 kg, it may have been

143 possible for a small group of foragers or even a single hunter to transport entire carcasses back to the site. The abundance of skeletal parts in the Pacbitun assemblages is compared here to different utility indices to determine if different currencies (e.g., meat, marrow, bone grease, raw material) guided transport decisions at Pacbitun.

In Chapter 6, it was argued that bone portions of low density were under- represented in the assemblages because of the effects of density-mediated processes, including differential preservation and carnivore ravaging. As a result, only elements of high bone density were selected for comparison with utility indices. According to the shape-adjusted reindeer bone density values provided by Lam and colleagues (1999), high-density portions for cervids are crania, mandibles, limb bone shafts, and the glenoid portion of the scapula. Although meat and marrow utility indices have been derived for white-tailed deer (Madrigal 2004), they were not used in this study because values were not available for all the elements which are of interest in this analysis. Three utility indices constructed for reindeer (Rangifer tarandus) were used instead (Binford 1978;

Metcalfe and Jones 1988; Morin 2007). Because the anatomical structure of ungulates is similar across species (Lam et al. 1999), using indices constructed for another cervid should provide comparable results.

The first utility index, named the simplified Meat Utility Index (MUI), measures the quantity of meat, marrow, and bone grease—or overall food content—of different skeletal parts (Metcalfe and Jones 1988). The simplified MUI is an intuitive measure of utility for the Pacbitun assemblages, because the residents of the site most likely consumed meat, as indicated by the presence of cutmarks, and extracted marrow from long bones (see section 7.8). The second utility measure can be termed the “grease index.” The data used to construct this index comes from an ethnographic episode

144 documented by Binford in spring 1971, during which he observed a Nunamiut woman selecting reindeer parts for bone grease rendering (Binford 1978). Data for phalanges in the MUI and grease index were obtained from Morin and Ready (2013), who corrected the values provided by Binford (1978) and Metcalfe and Jones (1988) to account for the fact that each foot in artiodactyls has two sets of three phalanges. The third economic measure is the Unsaturated Marrow Index (UMI) developed by Morin (2007). This index calculates the total quantity of unsaturated fatty acids in marrow-bearing bones, which are a major source of fats in ungulates.

Figure 7.2 compares the NNISP values for a restricted set of white-tailed deer elements to the reindeer MUI, grease index, and UMI. Because of small sample size, the early and late Middle Preclassic assemblages were combined in the analysis (Table 7.1).

It is expected that, if overall food value was the most important currency in transport decisions, the highest correlation should be found with the MUI. Instead, if within-bone fat content was the main currency in skeletal transport decisions, white-tailed deer skeletal abundances should correlate more strongly with the UMI or grease index.

All three correlations with the utility indices are weak and non-significant (Table

7.6). This result suggests that the Pacbitun Maya were relatively non-selective in terms of which long bones were transported to the site. This result may not be surprising. Given the small size of white-tailed deer in the tropics, carcasses of this animal could have been transported whole to the site by a small foraging party.

145

120

100 MAN MAX HUM

MC 60 FEM

%NNISP 40 TIB MT RAD 20 SCA 0 0 20 40 60 80 100 120 %MUI 120

100 HUM

MC 60 FEM

%NNISP TIB 40 RAD MT 20 SCA 0 0 20 40 60 80 100 120 %grease

120

100 HUM

MC 60 FEM

%NNISP TIB 40 RAD MT 20

0 0 20 40 60 80 100 120 %UMI

Figure 7.2 Comparison of white-tailed deer element frequencies (%NNISP) in the Pacbitun assemblages with the reindeer MUI, grease index, and UMI.

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Table 7.6 Spearman’s rank order correlations between skeletal part representation and the MUI, grease index, and UMI. n indicates the number of skeletal elements considered in the correlations.

Index n rs p MUI 9 0.08 0.85 Grease rendering 7 0.45 0.31 UMI 6 –0.61 0.20

Before accepting this interpretation of transport decisions at Pacbitun, it may be necessary to review factors that may confound the results. The absence of correlation may result from small sample size. Indeed, the sample of white-tailed deer elements selected for analysis has a NISP of 46, which is very small. Additionally, because only a small number of categories of elements were considered (MUI n = 9; grease index n = 7; UMI n

= 6), the statistical power of the comparisons is fairly low. An examination of the representation of other skeletal parts in the assemblages can provide further insight into transport decisions.

As mentioned in section 7.2, crania and mandibles are the most common elements in the assemblages, although they are only represented by teeth. Because teeth are more easily identified than bone fragments, their representation may be inflated in the samples.

Generally, it is considered that ungulate skulls are bulky, heavy, and of low nutritional value (Binford 1978). Ethnographic observations often report that skulls are discarded at kill sites because foragers do not want to invest energy in transporting skeletal elements which are considered of low profitability. However, as previously mentioned, this observation may not apply to small ungulates such as white-tailed deer which could be transported whole. At Pacbitun, the abundance of cranial and mandibular teeth suggests that these elements were frequently transported to the site.

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Feet are another bulky skeletal portion which may not be brought back to a central place because of its marginal utility value (Binford 1978). At Pacbitun, the first and second phalanges of white-tailed deer are quite abundant (phalanx 1: n= 9, %NNISP =

25.0; phalanx 2: n = 8, %NNISP = 22.2), whereas the third phalanx is only represented by three specimens (%NNISP = 8.3%). Five sesamoids are also present. This high frequency of feet elements (16.9%) may suggest that they were not frequently discarded at kill sites.

High abundances of these elements were also observed in the skeletal profiles of peccary

(46.2%) and red brocket deer (29.4%).

In sum, the absence of correlation between utility indices and long bone portions of high density suggests that the Pacbitun Maya regularly transported whole carcasses back to the site. The abundance of crania, mandibles, and feet elements in the Middle

Preclassic assemblages seems to support this interpretation.

7.8 Processing of skeletal parts

Given that the Pacbitun Maya were not selective regarding which skeletal elements should be transported back to the site, it may be possible to gain further insight into the decisions related to animal resource procurement by considering patterns of on- site skeletal part processing. At Pacbitun, long bones of white-tailed deer seem to have been systematically marrow-cracked. Green-bone fractures, which exhibit a curved fracture edge and a smooth fracture surface, were observed in high proportions (76.1%).

This type of fracture can be produced during marrow cracking, although trampling and carnivore ravaging may also create this type of damage. Although the sample is small, the presence of green-bone fractures on the long bones of smaller artiodactyls (peccary and brocket deer) suggests that these species were subjected to marrow exploitation as well.

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None of the notches in the Pacbitun assemblages were observed in association with gnaw marks or cutmarks. Although there is evidence for carnivore overprinting in the Pacbitun samples (Chapter 6), these animals only form a small fraction (3.8%) of the

Middle Preclassic assemblages. As a result, most notches are believed to have been produced by anthropogenic fracturing. It should be noted that heavy weathering of the material made it difficult to observe notches on fracture edges. Frequent abrasion of the edges may have obscured the identification of notches.

Intensity of marrow exploitation can be investigated through the examination of elements containing marginal amounts of marrow, such as the mandible, phalanges, calcaneum, and astragalus. In periods of food scarcity, humans may resort to extract marrow from some of these parts (Binford 1978). At Pacbitun, only white-tailed deer elements were considered in this analysis. In the early Middle Preclassic assemblage, all specimens of astragalus (n = 2), calcaneus (n = 3), first (n = 2) and second (n = 1) phalanges are fragmented, with the exception of two first phalanges which are complete.

None of these elements bear traces of carnivore gnawing. The picture is similar for the late Middle Preclassic sample. Most elements with a small marrow cavity are fragmentary

(calcaneus = 3/3; astragalus = 2/3; first phalanx = 5/5; second phalanx = 4/7). Although several specimens of calcaneum and first phalanges show edges consistent with green bone fractures, three specimens also display gnaw marks. As a result, it is difficult to determine whether phalanges and tarsals were fragmented as a result of marrow-cracking activities, carnivore ravaging, or both.

Bone grease rendering may be another indicator of nutritional stress. Indeed, the extraction of bone grease is a strenuous activity with high processing costs and low returns (Brink 1997). Grease rendering is the most destructive of all butchering processes

149 and can result in extensive fragmentation of long bone epiphyses. If this activity had been systematically practiced at Pacbitun, epiphyses would be severely depleted in comparison with long bone shafts. Although epiphyses of white-tailed deer are less common than shafts, differences in frequency (shafts = 53.2%; proximal = 27.4%; distal = 19.4%) between the bone portions are not sharp (K-S D = 0.29, p = 0.88). Morin (2012b:210) suggests that pounding and crushing marks associated with significant fragmentation of spongy bones may provide additional evidence for grease extraction. However, such marks were not observed in the Pacbitun samples. The destruction of epiphyses in the

Pacbitun assemblages more likely results from carnivore ravaging and post-depositional destruction (Chapter 6). Burning of spongy bone may also create a similar signature in the archaeological record (Morin 2010), but there is little evidence for the use of bone as fuel at Pacbitun given that none of the burned specimens (n = 38) are from cancellous bone.

Overall, marrow-cracking of long bones appear to have been systematic at the site.

It is not possible to ascertain if marginal marrow-bearing elements were also exploited due to a lack of data. There is also little evidence for grease rendering during the Middle

Preclassic at Pacbitun.

7.9 Pacbitun foraging strategies: A discussion

At Pacbitun, it appears that the Middle Preclassic Maya concentrated their efforts on some of the most profitable prey items, such as white-tailed deer and other smaller artiodactyls. However, less profitable prey types were also occasionally included in the diet breadth. These resources consist of a wide variety of small- and medium-sized game

(e.g., paca, agouti, opossum, rabbit, armadillo, iguana, and bird). It was suggested that felines and snakes were likely exploited by the Middle Preclassic Maya of Pacbitun.

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However, it is not possible to determine whether they were procured for food, as a source of raw material, and/or as a result of symbolic importance. Although there is evidence for the acquisition of exotic resources, such as marine fish, these resources seem to have been very marginal to the diet. Domestic dogs do not appear to have been an important source of food.

Vertebrate aquatic resources were also exploited, but in much smaller proportions than terrestrial animals. However, data presented by Stanchly (1999) regarding the exploitation of freshwater molluscs at Pacbitun tends to emphasize the importance of freshwater resources in the Maya diet. Freshwater molluscs represent 42.1% of the samples collected in 1995 and 1996. However, the importance of these animals may be over-estimated in this sample. First, molluscs are more easily identified than vertebrate remains. Second, as a result of their calcium carbonate composition, shell remains tend to preserve better than bone in highly acidic soil conditions. Third, although Stanchly argues that freshwater molluscs, particularly the jute, contributed significantly to the Preclassic diet, other authors (e.g., Solis 2011) disagree and conclude that their inclusion in the archaeological record may be incidental. For instance, 85.2% of the jute found during the

1995 and 1996 excavations were recovered from the thick secondary midden deposit which was laid on top of the Plaza B sub-structures at the end of the late Middle

Preclassic period. It is possible that the jute were included in the midden as part of riverbed soils which would have been used to level the plaza floor. Overall, these data suggest that freshwater resources were infrequently acquired at Pacbitun. Because they are slow-moving animals which can be easily captured, turtles probably constituted an exception to this pattern.

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Cannon (2003) explains that, when high-ranked resources are depleted from an environment, foragers need to invest more time in pursuing and acquiring lower-ranked prey types in order to continue maximizing their net delivery rate. This seems to have been the case at Pacbitun. Indeed, the scarcity of tapirs, the highest-ranked prey type, in the Pacbitun assemblages suggests that this prey item was infrequently encountered, possibly as a result of resource depression. Although the Pacbitun Maya are believed to be among the first settlers of the southern rim of the Belize River Valley, animal populations may have been under hunting pressure during earlier time periods, such as the

Early Preclassic (2000–1000 BC) or even the Archaic period. The Pacbitun Maya seem to have responded to this situation by concentrating their foraging efforts on a relatively narrow set of smaller-sized, but still highly profitable, prey items.

In sum, white-tailed deer, a relatively high-ranked species, appear to have formed the bulk of the diet at Pacbitun. Lower-ranked artiodactyls were also included in the diet, whereas a variety of other items may have been taken on occasion. The high frequency of long bones of white-tailed deer in the assemblages, combined with the abundance of crania, mandibles, tarsals, and phalanges, points to the frequent transport of whole carcasses. Animals were presumably exploited for their meat and marrow. Foraging clearly focused on the exploitation of terrestrial habitats, particularly those located near the site, with occasional trips taken to distant locales, such as mature forests and water sources. Although the early and late Middle Preclassic samples sometimes needed to be combined, the patterns of both assemblages seem consistent in terms of diet breadth, habitat use, and skeletal element exploitation.

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7.10 Subsistence strategies in the southern Maya lowlands

As mentioned earlier, research on the subsistence of the Maya during the Middle

Preclassic period is very limited. Only a handful of faunal assemblages recovered in the southern lowlands and dating to this time period have been analyzed (Table 2.2).

Unfortunately, some of these assemblages are affected by small sample size (e.g., Altar de Sacrificios, Seibal), which considerably limits interpretation of past subsistence strategies. Another prevalent problem is the lack of taphonomic data, as information on recovery methods and taphonomic history is not always available. Moreover, although evidence for anthropic manipulation is often mentioned, other factors that may cause differential preservation of faunal remains, including carnivore ravaging, weathering, and other density-mediated processes, are not reported. Keeping in mind these issues, one site from each of the sub-regions defined in Chapter 2 (i.e., Petén region of Guatemala, Belize

River Valley, and northern Belize) was selected for comparison with the Pacbitun assemblages. It should be noted that questions regarding the exploitation of meat, marrow, or fat from skeletal elements of large mammals could not be explored because this issue is rarely addressed by the studies.

The site of Cahal Pech is the only Middle Preclassic site from the Belize River

Valley for which we have faunal data. Cahal Pech is located on the banks of the Belize

River (Figure 2.1) and was occupied as early as the end of the Early Preclassic period

(1200–900 BC). The faunal dataset presented in this analysis was recovered exclusively from the Tolok group, a small community established in the periphery of the site core

(Powis et al. 1999). All faunal remains were found in an undisturbed midden deposit dated to the latter phase of the late Middle Preclassic period (450–300 BC).

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The Cahal Pech assemblage is dominated by fish remains (95.4%, Table 7.7), the majority of which have not been identified beyond class level (Powis et al. 1999). This pattern is not surprising given that Cahal Pech is located at the junction of three major river systems, the Macal, Mopan, and Belize Rivers. Marine fish (0.9%) have also been identified. Their presence at an inland site located some 110 km away from the Caribbean coast is attributed either to the existence of long-distance trade networks or to direct exploitation of the coast by the Cahal Pech Maya. Interestingly, the presence of skull elements from marine fish, some of which present traces of burning, suggests that marine resources were transported whole to the site where they would have been processed and consumed (Powis et al. 1999). Terrestrial mammals are the second most important taxonomic group in the sample (2.0%), but they are only represented by eight taxa, including white-tailed deer, brocket deer, domestic dog, paca, armadillo, rabbit, opossum, and small rodents. White-tailed deer is the most common mammal at Cahal Pech (0.7% of total NISP), whereas other artiodactyls are rare (<0.1%). Powis and colleagues (1999) suggest that most of these species were acquired within the local environment. They conclude that the foraging strategies at Cahal Pech would have focused on the procurement of terrestrial herbivores, marine reef fish, and small quantities of freshwater fish. It should be noted that preliminary analyses of faunal assemblages recovered from

Structure B-4 at Cahal Pech suggests that the residents of the site core focused on the exploitation of terrestrial mammals (57.7%) (Stanchly and Dale 1992). Fish was rarely identified in these last assemblages (6.5%). One wonders whether this disparity in taxonomic composition results from variations in recovery procedures, differential preservation, spatial distribution of faunal remains, or foraging strategies.

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Table 7.7 Percentages of vertebrate taxa identified in Middle Preclassic assemblages at Pacbitun (this study), Tolok group at Cahal Pech (Powis et al. 1999), Colha (Shaw 1999) and Bayak (Emery 2010).

Pacbitun Cahal Pech Colha Bayak Scientific Name Common Name n = 292 n = 2171 n = 1550 n = 268 Osteichthyes 2.7 95.4 40.8 38.9 Siluriformes Catfish 0.3 0.1 2.3 Serranidae Grouper 0.3 <0.1 0.8 Cichlidae Cichlid 5.3 Lepisoteiformes Gar Sparisoma spp. Parrotfish 0.3 0.7 Lachnolaimus spp. Hogfish <0.1 Lutjanidae Snappers <0.1 Unidentified fish 1.7 94.4 40.8 30.5 Amphibia 0.3 0.0 0.0 0.0 Rhinella marina Marine toad 0.3 Reptilia 10.6 0.8 47.5 53.1 Iguanidae Iguana 1.7 0.1 Testudines Turtle 5.5 0.1 17.5 32.4 Dermatemys sp. Central American River turtle 0.8 5.0 Emydidae Pond turtles 10.3 Trachemys scripta Terrapin 4.1 6.1 Rhinoclemmys areolata Furrowed wood turtle 0.3 Staurotypus sp. Musk turtles 3.1 Chelydra serpentina Common snapping turtle 0.4 6.1 Kinosternon spp. Mud turtle 0.3 11.7 Colubridae Colubrid 2.1 0.3 Viperidae Viper 1.0 0.5 Crocodylidae Crocodile 0.4 Unidentified reptile 0.6 1.7 Aves 1.7 1.7 0.4 0.8 Galliformes Turkey, curassow 0.3 0.1 Unidentified bird 1.4 1.6 0.4 0.8 Mammalia 84.6 2.0 11.2 7.3 Didelphis spp. Common opossum 1.4 0.4 0.5 Dasypus novemcinctus Nine-banded armadillo 12.0 0.1 0.7 Canidae Dog, fox 0.3 0.8 Canis lupus familiaris Domestic dog 0.7 0.1 2.2 Nasua narica White-nosed coati 0.3 Mustela frenata Long-tailed weasel 0.3 0.1 Felidae Cats 2.1 Tapirus bairdii Tapir 0.3 Artiodactyla Artiodactyl 0.7 0.4 Tayassuidae Peccary 4.5 0.1 0.4 Cervidae Cervid 2.4 1.7 Odocoileus virginianus White-tailed deer 47.6 0.7 4.4 5.0 Mazama americana Red brocket deer 5.8 <0.1 0.1 Rodentia 0.2 0.8 Orthogeomys spp. Pocket gopher 2.4 0.3 Dasyproctidae Agoutis, pacas 0.3 0.1 0.4 Cuniculus paca Paca, gibnut 1.0 0.3 Dasyprocta punctata Agouti 1.0 0.2 0.4 Sylvilagus spp. Rabbit 1.4 0.1 0.1 Total 100.0 100.0 100.0 100.0

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A group of three sites in northern Belize, Colha, Cuello, and K’axob, was presented in Chapter 2 (Table 2.2). Because taxonomic composition is similar at the three sites (Kruskall-Wallis ANOVA4 H = 3.71, p = 0.155), only one site is presented here.

Colha is located in northern Belize, roughly 45 km from the Caribbean Coast, near

Rancho Creek (Shaw 1999). The faunal assemblages discussed here were recovered from middens associated with domestic structures (Operations 2012 and 2031). During the

Middle Preclassic, the Colha Maya appear to have taken advantage of local terrestrial and aquatic habitats and strongly focused on the exploitation of wetlands. This is evidenced by the predominance of fish (40.8%) and turtles (45.2%) in the samples (Table 7.7).

Unfortunately, although both freshwater and marine fish were identified, the majority of fish remains could rarely be identified beyond class level. According to Shaw (1999), the small size of the fish vertebrae is more consistent with the anatomy of freshwater than marine fish. Therefore, she believes that most fish in the samples are from freshwater habitats. A fuller analysis of the fish remains will be needed to resolve this issue. A variety of terrestrial mammals (n = 11) were also exploited and form a small portion of the assemblages (11.2%). White-tailed deer (4.4%) and the domestic dog (2.2%) are the most common mammals.

From the three sites in the Petén region of Guatemala, only Bayak (Emery 1997) had a sample size sufficient for comparisons with other sites. Indeed, the identified vertebrate assemblages from Altar de Sacrificios (n = 24) and Seibal (n = 72) are very small. Bayak is a very small Preclassic site located on the edge of the Petexbatun Lake, in

Guatemala. Fish (38.9%) and turtles (52.7%) form the bulk of the samples, whereas

4 The Kruskall-Wallis ANOVA test was preferred over the chi-square test of independence because the assumptions of the latter test were not met by the datasets compared in this analysis.

156 terrestrial mammals, dominated by white-tailed deer (5.0%), only constitute a small part of the assemblage (7.3%). Fish are represented by almost equal proportions of cranial and axial elements, suggesting that they were used whole or prepared at the site. Analysis of habitat fidelity for the site of Bayak shows that riverine (75.5%) and swamp (21.8%) habitats were frequently used by the residents of the site (Emery 1997). These results are consistent with the location of the site on an elevated shoreline of the Petexbatun River.

Non-local faunal remain, such as marine fish, were not identified at this inland site.

Overall, data from Pacbitun, Colha, Cahal Pech, and Bayak suggest that the

Middle Preclassic Maya adopted foraging strategies consistent with the habitats in which they lived. Regional variation in species use is relatively high. In fact, differences in taxonomic composition between the four faunal assemblages are statistically significant

(Kruskal-Wallis ANOVA H = 16.8, p = 0.0008) (Table 7.8). Kolmogorov-Smirnov tests show that differences are not statistically significant for two pairs of sites: Cahal Pech and

Colha, and Cahal Pech and Bayak, possibly as a result of similar strategies of animal procurement. Indeed, faunal analyses show that the residents of these sites largely subsisted on fish and did not exploit terrestrial mammals in large quantities.

Table 7.8 Results of the Kolmogorov-Smirnov tests for the Middle Preclassic assemblages from the southern lowlands.

Sites Pacbitun Cahal Pech Colha Bayak Pacbitun – <0.001 <0.001 <0.001 Cahal Pech 0.75 – 0.31 0.14 Colha 0.60 0.33 – 0.002 Bayak 0.67 0.44 0.55 – Values below the dashes correspond to the D value, while those above are the p-values.

During the Middle Preclassic, the emphasis appears to have been placed on the exploitation of resources available in local micro-environments. Pacbitun, a site

157 surrounded by the tropical forest, is the only one where the population apparently focused on the exploitation of terrestrial animals of intermediate and large size. Conversely, the

Maya of Colha, which cultivated both milpas and wetlands, took advantage of the resources they could acquire in the vicinity of agricultural fields. This includes turtles, freshwater fish and, to a lesser extent, terrestrial mammals. The residents of Bayak, which lived on the shore of a large lake, focused nearly exclusively on the exploitation of freshwater fish and turtles. Cahal Pech is the only site which presented variation in the taxonomic composition of the assemblages recovered at the site. At Tolok, fish clearly dominate the sample, whereas preliminary analyses of samples recovered from the site core (Structure B-4) indicate that terrestrial mammals were the most common resources acquired in this part of the site. Because of a lack of data on the taphonomic history of these assemblages and the preliminary nature of the faunal analysis of Structure B-4, it is not possible to identify the cause(s) of this variation.

The presence of marine fish at some of these sites is also surprising. Table 7.9 presents the frequencies of marine fish at southern lowlands sites, with the average distance between each site and the Caribbean coast. As expected, no specimens of marine fish were recovered at the sites located in the Petén region of Guatemala, at a distance of

250–400 km from the coast. However, it is surprising that remains of marine fish were not identified at the two sites located closest to the Caribbean Sea, that is, K’axob and

Colha. It is suggested that this absence may result from an incomplete analysis of the fish samples. Indeed, significant quantities of fish remains were recovered from both sites, but a majority of specimens were not identified beyond class level. This explanation remains speculative. Small quantities of marine fish were also identified at Cuello, a site located about 35–50 km from the coast.

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Table 7.9 Abundances of marine fish at southern lowlands Maya sites during the Middle Preclassic period, with average distances from the coast of the Caribbean Sea.

Sites NISP % Distance K'axob 0 0 15–20 km Colha 0 0 20 km Cuello 30 0.49 35–50 km Cahal Pech 28 1.18 110 km Pacbitun 2 0.68 150 km Bayak 0 0 250–300 km Seibal 0 0 300 km Altar de Sacrificios 0 0 400 km Sources: Altar de Sacrificios (Pohl 1990), Bayak (Emery 2010), Cahal Pech (Stanchly 1995; Powis et al. 1999), Cuello (Wing and Scudder 1991; Carr and Fradkin 2008), Colha (Shaw 1999), K’axob (Masson 2004a), Pacbitun (this study), and Seibal (Pohl 1990).

The most surprising finds come from the inland sites of Cahal Pech and Pacbitun, where remains of parrotfish, grouper, and snapper were identified. Stable isotope analysis of carbon and nitrogen at Cahal Pech suggests that reef fish were consumed on a regular basis. This result appears incompatible with the location of the site, as well as with the very small amounts of marine fish in the archaeological assemblages (1.2%). It is suggested that differential preservation, sampling strategy, and/or difficulty of distinguishing between freshwater and marine fish species may explain the scarcity of marine fish at Cahal Pech, but also at other southern lowland sites. Overall, marine fish only formed a very small portion of the assemblages at the sites where they have been identified. No correlation was found between the quantity of fish identified in the faunal assemblages and a site’s distance from the Caribbean Sea (rs = –0.13, p = 0.74).

Additionally, it was not also possible to determine whether the presence of marine fish at

Middle Preclassic sites reflect long-distance fishing trips or the beginnings of exchange of goods between communities located in different habitats.

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A point should be made concerning the argument that domestic dogs were heavily used as a source of food during the Preclassic period. This pattern, which was first inferred at Cuello (Clutton-Brock and Hammond 1994), is not supported at Pacbitun,

Bayak, or Cahal Pech. However, Shaw (1991) reported that there is unambiguous evidence for the consumption of dogs at Colha in the form of cutmarks. She also explains that many bones are burned, a situation which she attributes to the cooking of these animals for food consumption. It is suggested here that the use of dogs as a source of food during the Middle Preclassic period may have been a restricted to northern Belize.

Lastly, the taxonomic composition from Colha, Cahal Pech, and Bayak suggests that, contrary to what was found at Pacbitun, the prey items most frequently taken were not always be the largest in terms of body mass. In fact, terrestrial mammals only form a small portion of the assemblages, with ungulates generally constituting the most abundant terrestrial taxa (Cahal Pech = 0.7% of NISP; Colha = 6.3%; Bayak = 6.5%). According to the central place forager prey choice model, the scarcity of large terrestrial game may indicate depletion of these resources in patches located near the three sites. In fact, the emphasis placed on the exploitation of other less profitable resources at Cahal Pech,

Bayak, and Colha may reflect a broadening of the diet. Fish, an important resource at the three sites, was perhaps favored because it can be mass-collected with the use of nets.

This fishing technology can increase the net return rate of energy acquisition, particularly if small fish do not need to be processed extensively. The exploitation of turtles also seems to have been the focus of foraging strategies at Bayak and Colha, two sites which had access to many water sources. As previously mentioned, turtles are slow-moving animals which can provide important quantities of meat for little costs. As such, they may always have been part of the optimal diet.

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CHAPTER 8: CONCLUSION

This study has investigated the foraging strategies related to the acquisition of animal resources during the Middle Preclassic period at the ancient Maya site of Pacbitun.

This concluding chapter revisits the objectives outlined in Chapter 1 and summarizes the findings of the faunal analysis. It also examines several limitations of this study and considers the significance of this research. Recommendations for future research are also provided.

8.1 Research summary

The results of this study are summarized here according to the objectives outlined in Chapter 1.

1) Taphonomic analysis:

A detailed analysis of the Pacbitun faunal assemblages has revealed that several taphonomic processes have shaped the samples. The use of two different recovery methods does not seem to have affected the taxonomic composition of the assemblages, although selective recovery may have depleted the assemblages of specimens smaller than 1 cm. A majority of faunal remains have been severely affected by weathering, a situation which impeded taxonomic identification and possibly obliterated the identification of marks left on bone surfaces. Carnivores, in particular domestic dogs, appear to have had secondary access to the bone deposits and possibly altered the assemblages. Depletion of elements and bone portions of low density may also result from post-depositional destruction and perhaps burning. Findings of the taphonomic analysis were similar for both the early and late Middle Preclassic assemblages, although bones surfaces were more poorly preserved in the late Middle Preclassic sample.

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2) Analysis of foraging strategies at Pacbitun:

Archaeozoological analysis of the faunal samples indicates that the Middle

Preclassic Maya of Pacbitun exploited a wide variety of vertebrate animal resources, including mammals, reptiles, fish, and birds. Animal procurement strategies focused on the exploitation of ungulate species, with white-tailed deer constituting by far the primary prey item. The most profitable prey types according to the body mass scale, that is, the tapir, jaguar, and puma, seem to have been infrequently encountered. Low-ranked prey taxa were occasionally exploited. These included a variety of small- and medium-sized game. Secondary growth forests and agricultural fields, two habitat types which would have been located at short distance from the site, were the primary focus of hunting activity. The low abundances of freshwater resources and forest-dwelling animals suggest that riverine areas and mature forests were less heavily utilized. Analysis of skeletal patterns suggests that the carcasses of large ungulates were transported whole to the site.

Animal resources appear to have been exploited for their meat and marrow fat content.

Labor-intensive activities, such as bone grease rendering and marrow-cracking of marginal marrow-bearing elements, do not appear to have been practiced on a regular basis.

3) Animal resource exploitation in the southern Maya lowlands:

Comparisons of faunal assemblages from four Maya sites suggest that, during the

Middle Preclassic, procurement of animal resources focused on taxa which could be acquired in local habitats. Exotic resources, such as marine fish, were also imported in small quantities to sites located inland. This possibly attests to the presence of long- distance trade networks during this early time period. The foraging patterns identified at

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Pacbitun differ from those observed at Cahal Pech, Colha, or Bayak. People at these three sites relied more heavily on fish and, in the case of Colha and Bayak, turtles. While large terrestrial game is the prey item most frequently taken at Pacbitun, it only forms a fraction of the assemblages at other Middle Preclassic sites. It is suggested that depletion of large- sized prey types in patches located near the sites may have led to a broadening of the diet and the subsequent exploitation of less profitable resources, such as fish.

8.2 Limitations and significance

The primary limiting factor of this investigation is the small sample size of the

Pacbitun assemblages (early Middle Preclassic NISP = 125; late Middle Preclassic NISP

= 167). In some instances, this situation precluded comparisons of the early and late

Middle Preclassic samples. Additionally, because of small sample size, many species were only represented by one or two specimens. In those cases, the question remains of whether these taxa are rare in the assemblages as a result of sample size or because they were not exploited by the ancient Maya.

Additionally, as a result of small sample size, the statistical power of this study is fairly low. Indeed, in order for statistical tests to be robust (i.e., to identify significant relationships among data), samples must be relatively large. This is because the effect of outliers is much greater in small samples, a situation which can obscure or exaggerate trends within the sample. To control for this situation, nonparametric statistical tests were employed in this study because they are less prone to the effects of small sample sizes and abnormally distributed data than parametric tests (Chenorkian 1996). Despite small sample size, trends emerged from the analysis of the Pacbitun faunal data, such as a clear focus on the exploitation of white-tailed deer by the residents of the site. Hopefully,

163 continuing research at Pacbitun should help to increase the size of the faunal samples and confirm the patterns observed in this study.

Despite these limitations, this study shows that the Pacbitun Maya took advantage of resources locally available, focusing mainly on large prey items such as artiodactyls.

The data also lend support to the notion that foraging strategies during the Middle

Preclassic were highly focused on the exploitation of local habitats. Although there is evidence for the acquisition of animal resources from distant locales, the vast majority of prey items were acquired in the vicinity of the sites.

8.3 Future Directions

Although it was shown in this study that the use of different screen mesh sizes did not affect the Pacbitun assemblages in terms of fragment size or taxonomic composition, it is suggested that finer mesh screens should be used in the future in order to increase the chances of recovering smaller specimens and/or taxa. However, the use of finer mesh screens is a relatively time-consuming activity, particularly when soils are of clay composition as it is the case at Pacbitun. A reasonable strategy might be to draw random soil samples and to screen them with fine mesh screens (e.g., 1/8 or 1/16 inch mesh screen) to determine which mesh size is the most adequate for maximizing the recovery of faunal remains. It is should be noted that the use of finer mesh screens do not necessarily lead to the increased recovery of faunal remains because small remains may have disappeared in poorly preserved faunal assemblages.

Because the Maya exploited animals for economic, social, political, and religions purposes, faunal data alone may not always be sufficient to detect the multiple ways in which animals were used. As a result, it is suggested that many lines of evidence should

164 be used to interpret faunal data. Alongside iconographic, ethnohistoric and ethnographic records which are typically used by Maya zooarchaeologists, evidence for past animal use may be obtained from more recently developed methods, such as residue analysis and stable isotope analysis. For instance, at Pacbitun, the analysis of habitat fidelity only provided coarse information about the provenience of the animals exploited by the site’s residents. However, given the location of Pacbitun between two ecozones, the tropical rainforest and the pine ridge, it may be interesting to determine if prey taxa were preferably acquired from one of these two ecozones or even if animals, in particular terrestrial species, might have been procured from the Belize River Valley or other regions in the southern lowlands. Had the scope of this research been larger, this question could have been investigated through strontium isotope analysis.

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APPENDIX A

Pacbitun faunal datasets by time period

201

ABBREVIATIONS

Identification

Code Scientific name Common name

ART Artiodactyla Artiodactyls CER Cervidae Cervids ODO Odocoileus virginianus White-tailed deer MAZ Mazama americana Red brocket deer TAY Tayassuidae Peccaries TAP Tapirus bairdii Baird’s tapir CAN Canidae Canids DOG Canis lupus familiaris Domestic dog COA Nasua narica Coatimundi MUS Mustela frenata Weasel ARM Dasypus novemcinctus Armadillo OPOV Didelphis virginiana Virginia opossum OPOM Didelphis marsupialis Common opossum CAT Felidae Cats JAG Panthera onca Jaguar PUM Puma concolor Puma OCE Leopardus pardalis Ocelot MAR Leopardus wiedii Margay GOP Orthogeomys spp. Pocket gophers DAS Dasyproctidae Agouti, paca PAC Cuniculus paca Paca AGO Dasyprocta punctata Agouti RAB Sylvilagus spp. Rabbits AVES Aves Birds GAL Galliformes Turkey, guan COL Colubridae Colubrid VIP Viperidae Vipers IGU Iguanidae Iguanas TES Testudines Turtles KIN Kinosternon spp. Mud turtle FISH Unidentified fish ICT Ictaluridae Catfish SER Serranidae Grouper PAF Sparisoma spp. Parrotfish BUF Rhinella marinus Marine toad STM Small terrestrial mammal MTM Medium terrestrial mammal LTM Large terrestrial mammal UNIM Indeterminate mammal UNI Indeterminate specimen

202

Bone element

ANT Antler fragment ILM Ilium SKL Skull fragment INN Innominate fragment FRN Frontal ISH Ischium OCC Occipital PUB Pubis NAS Nasal PAT Patella PAR Parietal FEM Femur PET Petrous (bulla) TIB Tibia ROS Rostrum FIB Fibula TEM Temporal TFB Tibia-fibula ZYG Zygomatic AST Astragalus PMX Premaxilla CAL Calcaneus MAN Mandible CUB Cuboid MAX Maxilla GC Greater cuneiform IN Incisor MAL Malleolar CN Canine NAV Navicular PM Premolar SC Smaller cuneiform MO Molar NVB Naviculo-cuboid DP Deciduous premolar SES Sesamoid TTH Tooth fragment MTC Metacarpal HYD Hyoid MTP Metapodial MTT Metatarsal ATL Atlas PH1 Phalanx 1 AXI Axis PH2 Phalanx 2 CEV Cervical vertebra PH3 Phalanx 3 TRV Thoracic vertebra PHA Phalanx fragment LMV Lumbar vertebra SHL Shell CDV Caudal vertebra DRS Dorsal spine VER Vertebral fragment LBF Long bone fragment SAC Sacrum UNI Unidentified bone element STX Sternum RIB Rib SYN Synsacrum CC Coastal cartilage TBT Tibiotarsus CMT Carpometacarpus SCA Scapula TMT Tarsometatarsus HUM Humerus PPX Proximal phalanx (wing) RAD Radius DPX Distal phalanx (wing) RUL Radius-ulna UNG Ungus (talon) ULN Ulna RNG Tracheal ring CAP Capitatum FUR Furcula HAM Hamatum COR Coracoid LUN Lunate LSA Lumbosacral PIS Pisiform QUA Quadrate SCP Scaphoid SCL Scapholunar TRQ Triquetrum CUN Cuneiform

203

End PE Proximal end PS Proximal shaft MS Middle shaft DS Distal shaft DE Distal end ANT Anterior POST Posterior CRA Cranial CAU Caudal UP Upper tooth LO Lower tooth ACE Acetabulum CEN Centrum (vertebra) W Whole W-S Fish vertebra with intact centrum but lacking all spines S Shaft fragment F Fragment

Side L Left R Right AX Axial element

Fusion FU Fused U Unfused I Intermediate (fused by line clearly visible)

Only for deer teeth FAW Fawn: <1 year old YEA Yearling: 1–2 years old ADU Adult: >2 years old

Max length <1 measuring less than 1 cm 1–2 measuring between 1.00–1.99 cm 2–3 measuring between 2.00–2.99 cm 3–4 measuring between 3.00–3.99 cm 4–5 measuring between 4.00–4.99 cm

Surface state PO Poor DA Damaged SD Slightly damaged IT Intact

204

Taphonomy CRK Cracking RT Root EX Exfoliation SH Sheeting STG Staining CT Cut mark PN Percussion notch DR Drilled BUR Burned GN Gnawing

Table A.1 Pacbitun dataset for the early Middle Preclassic period

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-08BO-014-01 B 1 6 UNIM UNI F 3 <1 DA EX

PAC-08BO-014-02 B 1 6 UNIM UNI F 5 1–2 DA EX

PAC-08BO-015-01 B 1 6 UNIM UNI F 1 1–2 SD EX

PAC-08BO-015-02 B 1 6 LTM LBF F 1 5.5 1.9 DA EX

PAC-08BO-015-03 B 1 6 DAS IN F 1 1.6 0.4 DA EX

PAC-08BO-015-04 B 1 6 IGU ILM F L 1 U 1.3 1.3 DA EX, RT PAC-08BO-020-01 B 1 7 LTM VER F AX 1 FU 2.1 1.5 SD EX, GN PAC-08BO-020-02 B 1 7 TAY RAD PS+MS+DS R 1 7.3 1.8 SD EX, RT

PAC-08BO-022-02 B 2 6 ODO RIB POST R 1 2.6 1.2 SD EX, RT

PAC-08BO-024-01 B 1 7 UNI UNI F 2 <1 PO EX

PAC-08BO-025-01 B 1 7 LTM CDV F AX 1 2.1 1.0 DA EX, GN

PAC-08BO-026-01 B 2 7 UNI UNI F 1 <1 PO EX

PAC-08BO-026-02 B 2 7 UNIM LBF F 1 2.8 1.1 PO EX, RT

PAC-09BO-007/008-01 B 1 6B UNI UNI F 1 3–4 DA EX, RT

PAC-09BO-007/008-02 B 1 6B LTM LBF F 1 4.8 1.1 PO EX, RT

PAC-09BO-007/008-03 B 1 6B ODO ULN PE+PS R 1 FU 8.0 3.2 SD EX, RT, GN PAC-09BO-007-01 B 1 6B UNI UNI F 3 1–2 DA EX

PAC-09BO-007-02 B 1 6B UNIM LBF F 1 2.6 0.8 SD EX

PAC-09BO-007-03 B 1 6B LTM LBF F 1 4.1 1.1 PO EX, SH, STG

PAC-09BO-007-04 B 1 6B ARM MTT 4 W R 1 FU 1.7 0.7 DA EX, RT PAC-09BO-007-05 B 1 6B JAG PH2 W L 1 FU 2.0 1.4 DA EX, CT PAC-09BO-007-06 B 1 6B MTM-LTM SCA PRO L 1 FU 1.7 1.3 PO EX, GN PAC-09BO-008-01 B 1 6B UNI UNI F 2 <1 PO EX

PAC-09BO-008-02 B 1 6B UNIM UNI F 5 1–2 PO EX

PAC-09BO-008-03 B 1 6B UNIM LBF F 2 2–3 DA EX

205

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-09BO-008-04 B 1 6B ODO FEM DE R 1 U 2.6 2.5 SD EX PAC-09BO-008-05 B 1 6B ODO MO UP 1 1.4 0.9 DA EX

PAC-09BO-008-06 B 1 6B ARM TIB PE+PS L 1 U 1.6 1.2 PO EX PAC-09BO-008-07 B 1 6B ARM TIB PE+PS+MS R 1 U 3.2 1.5 SD EX, GN PAC-09BO-008-08 B 1 6B ARM FIB MS+DS L 1 2.4 0.5 SD EX, STG

PAC-09BO-008-09 B 1 6B ART MO-PM F 1 1.6 1.0 DA EX

PAC-09BO-008-10 B 1 6B MTM-LTM UNI F 1 2.7 1.3 DA EX, CT

PAC-09BO-008-11 B 1 6B OPOV MAN DIS L 1 FU 1.6 1.0 DA EX, RT PAC-09BO-009-01 B 1 6B ODO MO1-2 UP R 1 1.4 1.3 DA EX, CRK

PAC-09BO-010-01 B 1 6B UNI UNI F 7 <1 PO EX

PAC-09BO-010-02 B 1 6B UNIM UNI F 10 1–2 SD EX

PAC-09BO-010-03 B 1 6B UNIM UNI F 15 1–2 PO EX

PAC-09BO-010-04 B 1 6B UNIM UNI F 3 2–3 PO EX, CRK

PAC-09BO-010-05 B 1 6B UNIM LBF F 1 2.0 0.7 SD RT, BUR

PAC-09BO-010-06 B 1 6B UNIM LBF F 1 1.6 0.9 IT RT

PAC-09BO-010-07 B 1 6B STM LBF F 1 2.4 0.5 PO EX, RT

PAC-09BO-010-09 B 1 6B LTM UNI F 1 3.8 1.5 PO SH, CRK

PAC-09BO-010-10 B 1 6B STM-MTM SKL F 1 1.1 0.9 SD

PAC-09BO-010-11 B 1 6B ODO AST PRO R 1 FU 2.7 1.5 DA EX PAC-09BO-010-12 B 1 6B ODO NVB F R 1 FU 2.2 1.2 SD EX PAC-09BO-010-13 B 1 6B MAZ CAP W L 1 FU 1.1 1.0 SD EX PAC-09BO-010-14 B 1 6B MTM-LTM MAN? F 1 2.9 1.3 PO EX, CRK, SH

PAC-09BO-010-15 B 1 6B ODO CAL PE+PS+MS L 1 FU 5.8 2.2 SD EX, CT PAC-09BO-010-16 B 1 6B ODO MTC PE+PS L 1 FU 3.1 1.5 SD EX, CRK PAC-09BO-010-17 B 1 6B ODO AST DE R 1 FU 1.8 1.7 SD EX PAC-09BO-010-18 B 1 6B ODO RIB POST L 1 FU 2.2 1.0 SD EX PAC-09BO-010-19 B 1 6B ODO CAL DE L 1 FU 2.8 1.4 SD EX

206

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-09BO-010-20 B 1 6B ODO PH1 W 1 FU 3.9 1.4 SD EX

PAC-09BO-010-21 B 1 6B TAY AST F R 1 FU 1.4 1.0 DA EX, GN PAC-09BO-010-22 B 1 6B ARM TFB DE R 1 U 2.2 0.9 PO EX, RT, CRK PAC-09BO-010-23 B 1 6B ARM TIB DS+MS L 1 2.9 1.0 DA EX, RT

PAC-09BO-010-24 B 1 6B LTM VER F AX 1 1.8 1.3 DA EX

PAC-09BO-018-01 B 2 6 UNIM UNI F 3 1–2 PO EX

PAC-09BO-018-02 B 2 6 UNIM UNI F 1 1–2 SD EX

PAC-09BO-018-03 B 2 6 CER ULN DS R 1 1.3 0.6 DA EX

PAC-09BO-018-04 B 2 6 ODO TIB MS R 1 3.2 1.3 SD EX, STG

PAC-09BO-018-05 B 2 6 ART PHA DS+DE 1 1.5 0.9 SD

PAC-09BO-018-06 B 2 6 ODO TIB PE L 1 FU 3.4 2.8 SD EX, GN PAC-09BO-018-07 B 2 6 MTM ISH ACE L 1 FU 1.5 1.2 DA EX, RT PAC-09BO-018-08 B 2 6 UNIM UNI F 1 2–3 PO EX, CRK

PAC-09BO-019/021 B 2 6B ODO FEM PS R 1 5.2 2.0 PO EX

PAC-09BO-019-01 B 2 6B MAZ ULN PS+MS R 1 5.5 0.7 DA EX, GN

PAC-09BO-019-02 B 2 6B UNI UNI F 1 2–3 PO EX, RT

PAC-09BO-020-01 B 2 6B UNI UNI F 5 <1 PO EX

PAC-09BO-020-02 B 2 6B UNIM UNI F 5 1–2 PO EX

PAC-09BO-020-03 B 2 6B UNIM UNI F 1 2–3 PO EX

PAC-09BO-020-04 B 2 6B UNIM LBF F 3 2–3 PO EX, RT, STG

PAC-09BO-020-05 B 2 6B UNIM LBF F 1 2–3 SD RT, STG

PAC-09BO-020-06 B 2 6B TAY SCA CRA R 1 FU 2.2 1.4 SD EX, RT PAC-09BO-020-07 B 2 6B LTM LBF F 1 5.1 1.7 PO SH, CRK

PAC-09BO-020-08 B 2 6B LTM UNI F 1 3.8 1.5 PO EX, RT

PAC-09BO-021-01 B 2 6B UNIM UNI F 2 1–2 PO EX

PAC-09BO-021-02 B 2 6B UNIM UNI F 1 1–2 DA EX

PAC-09BO-021-03 B 2 6B MAZ MTC PE+PS+MS+DS L 1 FU 8.1 1.3 SD EX, GN

207

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-09BO-021-04 B 2 6B AGO HUM MS+DS R 1 2.8 0.5 SD RT

PAC-09BO-021-05 B 2 6B TES SHL F 1 1.0 0.8 DA EX, STG

PAC-09BO-030-01 B 3 6A UNIM LBF F 1 1.9 0.7 DA EX, RT, STG

PAC-09BO-030-02 B 3 6A UNIM LBF F 1 2.5 0.9 DA EX, RT

PAC-09BO-030-03 B 3 6A UNI UNI F 1 <1 PO EX

PAC-09BO-031-01 B 3 6B AGO IN1 UP R 1 1.2 0.3 IT RT, STG

PAC-09BO-032-01 B 3 6B UNIM UNI F 1 1–2 PO EX, RT

PAC-09BO-041-01 B 4 6 UNIM UNI F 1 <1 SD EX

PAC-09BO-041-02 B 4 6 UNI UNI F 1 1–2 SD EX

PAC-09BO-041-03 B 4 6 UNIM UNI F 2 1–2 PO EX, RT

PAC-09BO-041-04 B 4 6 UNIM UNI F 1 2–3 PO EX, RT

PAC-09BO-041-05 B 4 6 TES SHL F 1 1.5 1.4 PO EX

PAC-09BO-041-06 B 4 6 TES SHL F 1 1.4 1.4 PO EX

PAC-09BO-045-01 B 4 6B UNI UNI F 1 1–2 PO EX

PAC-09BO-046-01 B 4 6B UNIM UNI F 1 1–2 PO EX

PAC-09BO-046-02 B 4 6B MTM-LTM LBF F 1 4.0 1.4 PO EX, RT, STG

PAC-09BO-047-01 B 4 6B UNI UNI F 6 <1 PO EX

PAC-09BO-047-02 B 4 6B UNI UNI F 5 <1 SD

PAC-09BO-047-03 B 4 6B UNIM UNI F 5 1–2 PO EX

PAC-09BO-047-04 B 4 6B UNIM UNI F 3 2–3 DA EX

PAC-09BO-047-05 B 4 6B UNIM UNI F 2 <1 DA EX

PAC-09BO-047-06 B 4 6B ODO ULN PE+PS L 1 FU 4.1 2.0 DA EX, RT, CRK, STG PAC-09BO-047-07 B 4 6B ODO CEV CRA AX 1 FU 4.8 4.3 DA EX, RT, CRK PAC-09BO-047-08 B 4 6B ODO FEM DE L 1 FU 2.2 2.1 PO EX, GN PAC-09BO-047-09 B 4 6B UNIM UNI F 3 1–2 DA EX, RT, CRK

PAC-09BO-047-10 B 4 6B UNIM UNI F 1 2–3 DA EX, RT

PAC-09BO-047-13 B 4 6B ODO SES W 1 FU 0.8 0.6 SD EX

208

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-09BO-047-14 B 4 6B ODO SES F 1 FU 1.2 1.0 SD

PAC-09BO-048-01 B 4 6B UNIM UNI F 1 <1 PO EX

PAC-09BO-048-02 B 4 6B UNIM UNI F 6 1–2 DA EX

PAC-09BO-048-03 B 4 6B UNIM UNI F 2 2–3 DA EX, RT

PAC-09BO-048-04 B 4 6B LTM UNI F 1 2.5 1.8 SD EX, RT

PAC-09BO-048-05 B 4 6B CAT CDV W AX 1 FU 1.3 0.5 PO EX, RT PAC-09BO-048-06 B 4 6B ODO PH1 PE 1 FU 1.6 0.9 PO EX

PAC-09BO-048-07 B 4 6B ODO GC W L 1 FU 1.6 0.9 PO EX PAC-09BO-048-08 B 4 6B LTM LBF F 1 6.0 1.9 DA EX, RT, CRK, STG

PAC-10BO-003-01 A 1 8 UNIM UNI F 1 1–2 DA EX, RT

PAC-10BO-008-01 A 1 6 COA TEM F R 1 2.4 1.8 PO EX

PAC-10BO-009-01 A 1 7 UNIM UNI F 1 1–2 PO EX

PAC-10BO-009-02 A 1 7 UNIM UNI F 1 1.9 0.8 SD EX, BUR

PAC-10BO-009-03 A 1 7 ODO ULN PE R 1 FU 6.6 3.1 PO EX, RT PAC-10BO-009-04 A 1 7 ODO RAD PE+PS L 1 FU 4.9 3.1 DA EX, RT PAC-10BO-009-05 A 1 7 UNIM UNI F 1 2–3

PAC-10BO-015-01 B 3 6B UNIM UNI F 1 1–2 PO EX

PAC-10BO-015-02 B 3 6B UNIM UNI F 1 1–2 SD EX

PAC-10BO-015-03 B 3 6B UNI UNI F 1 1–2 SD RT

PAC-10BO-016-01 B 3 6B STM-MTM LBF F 1 1.4 0.6 DA EX, RT, CRK

PAC-10BO-016-02 B 3 6B MAZ MTC PE+PS+MS R 1 FU 4.8 1.0 DA EX, RT, CRK, STG PAC-10BO-016-03 B 3 6B RAB FEM DE L 1 FU 0.8 0.8 SD EX PAC-10BO-016-04 B 3 6B MTM CN F 1 1.0 0.3 IT RT

PAC-10BO-016-05 B 3 6B GOP PM4 LO L 1 1.0 0.4 DA EX, RT

PAC-10BO-017-01 B 3 6B UNI UNI F 1 <1 SD EX

PAC-10BO-017-02 B 3 6B UNIM UNI F 3 1–2 PO EX

PAC-10BO-017-03 B 3 6B FISH VER W-S AX 1 0.5 0.4 SD

209

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-10BO-017-03 B 3 6B UNI UNI F 1 1–2 SD RT

PAC-10BO-018-01 B 3 6B UNIM UNI F 1 <1 PO EX

PAC-10BO-018-02 B 3 6B UNIM UNI F 1 1–2 PO EX

PAC-10BO-019-01 B 3 6B UNIM UNI F 1 <1 PO EX

PAC-10BO-019-02 B 3 6B UNIM UNI F 1 2–3 PO EX, RT

PAC-10BO-021/022 B 5 6 ODO MTC PE+PS+DS R 1 FU 7.5 1.8 PO EX, CRK, GN PAC-10BO-021-01 B 5 6 UNI UNI F 2 <1 PO EX

PAC-10BO-022-01 B 5 6 UNIM UNI F 5 <1 PO EX

PAC-10BO-022-02 B 5 6 UNIM UNI F 1 1–2 PO EX, RT

PAC-10BO-023-01 B 5 6 UNI UNI F 4 <1 PO EX

PAC-10BO-023-02 B 5 6 TES SHL F 1 2.2 1.3 PO EX, CRK

PAC-10BO-023-03 B 5 6 MAZ PH1 DE+DS+MS 1 FU 1.6 0.8 SD EX

PAC-10BO-024-01 B 5 6 UNIM UNI F 1 1–2 PO EX

PAC-10BO-024-02 B 5 6 PAC MO1 LO L 1 1.2 0.7 SD RT, STG

PAC-10BO-025-01 B 5 6B UNIM UNI F 5 <1 PO EX

PAC-10BO-025-02 B 5 6B UNIM UNI F 2 1–2 DA EX, RT

PAC-10BO-025-03 B 5 6B UNIM UNI F 1 3–4 DA EX, RT

PAC-10BO-025-04 B 5 6B ODO NVB W R 1 FU 2.8 2.4 DA EX, RT, CT PAC-10BO-025-05 B 5 6B OPOV ULN PE R 1 FU 1.1 0.6 SD EX, RT PAC-10BO-026-01 B 5 6B FISH VER W-S AX 1 0.5 0.4 IT

PAC-10BO-026-02 B 5 6B IGU MAN+TTH Mesial R 1 1.4 0.5 SD EX, RT

PAC-10BO-026-03 B 5 6B STM LBF F 1 0.8 0.3 SD EX

PAC-10BO-026-04 B 5 6B STM LBF F 1 2.1 0.4 SD EX

PAC-10BO-027-01 B 5 6B UNI UNI F 1 <1 SD

PAC-10BO-027-02 B 5 6B UNIM UNI F 1 1–2 PO EX, RT

PAC-10BO-028-01 B 5 7 UNIM UNI F 4 1–2 PO EX

PAC-10BO-028-02 B 5 7 UNIM UNI F 2 2–3 PO EX

210

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-10BO-028-03 B 5 7 ARM SHL F 1 1.0 0.6 DA EX

PAC-10BO-028-04 B 5 7 LTM RIB F 1 1.6 0.6 DA EX, STG

PAC-10BO-028-05 B 5 7 GOP TIB PS+MS R 1 1.3 0.5 PO EX, STG

PAC-10BO-028-06 B 5 7 UNIM UNI F 1 3–4 PO EX, RT

PAC-10BO-032-01 B 6 6 UNI UNI F 8 <1 PO EX

PAC-10BO-032-02 B 6 6 UNIM UNI F 6 1–2 PO EX

PAC-10BO-032-03 B 6 6 ARM SHL F 1 0.7 0.6 PO EX

PAC-10BO-032-04 B 6 6 GOP PM4 LO L 1 1.0 0.4 SD EX

PAC-10BO-033-01 B 6 6 UNI UNI F 1 <1 SD EX

PAC-10BO-033-02 B 6 6 UNIM UNI F 1 1–2 PO EX

PAC-10BO-033-03 B 6 6 STM UNI F 1 0.8 0.4 SD BUR

PAC-10BO-034A-01 B 6 6B UNI UNI F 7 <1 PO EX

PAC-10BO-034A-02 B 6 6B UNIM UNI F 1 1–2 PO EX, RT

PAC-10BO-034A-03 B 6 6B LTM LBF F 1 2.4 1.4 PO

PAC-10BO-034B-01 B 6 6 UNI UNI F 1 <1 PO EX

PAC-10BO-034B-02 B 6 6 UNIM UNI F 1 1–2 PO EX

PAC-10BO-035A-01 B 6 6B PAC MO3 LO R 1 1.7 0.7 SD RT

PAC-10BO-035B-01 B 6 6B UNI UNI F 2 <1 DA EX

PAC-10BO-035B-02 B 6 6B UNIM UNI F 6 1–2 PO EX

PAC-10BO-035B-03 B 6 6B UNIM LBF F 1 2.0 0.9 PO EX

PAC-10BO-035B-04 B 6 6B LTM LBF F 1 3.6 1.8 PO EX, RT

PAC-10BO-035B-08 B 6 6B STM LBF F 1 1.4 0.4 PO EX, STG

PAC-10BO-036A-01 B 6 6B UNI UNI F 2 <1 PO EX, STG

PAC-10BO-036B-01 B 6 7 UNI UNI F 4 <1 PO EX

PAC-10BO-036B-02 B 6 7 ODO LUN F R 1 1.7 0.9 SD EX

PAC-10BO-036B-03 B 6 7 ODO SCP F L 1 FU 1.6 0.8 PO EX PAC-10BO-036B-04 B 6 7 ARM SHL F 1 0.4 0.3 DA EX

211

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-10BO-038-01 B 8 6 UNI UNI F 2 <1 PO EX

PAC-10BO-038-02 B 8 6 ODO PH2 F 1 2.5 1.3 PO EX

PAC-10BO-040-01 B 10 6 UNI UNI F 3 <1 PO EX PAC-11BO-001-01 UNIM UNI F 1 2.8 1.2 A 1E 7 DA RT, BUR PAC-11BO-001-02 UNIM UNI F 1 1–2 A 1E 7 PO EX, RT PAC-11BO-002-01 A 1E 8 ODO PH1 DE+DS+MS 1 FU 2.5 1.1 SD EX, RT, STG

PAC-11BO-003-01 A 1F 7 MTM-LTM LBF F 1 2.7 1.3 PO EX, RT, BUR, DR UNIM UNI F 1 1.3 0.7 PAC-11BO-003-02 A 1F 7 SD RT, BUR UNI UNI F 1 1–2 PAC-11BO-003-03 A 1F 7 SD RT PAC-11BO-004-01 UNIM UNI F 1 A 1F 8 2–3 PO EX PAC-11BO-004-02 PH1 W 1 3.7 1.3 A 1F 8 ODO SD EX, CRK, BUR PAC-11BO-004-03 A 1F 8 ODO SCA PRO L 1 FU 8.4 4.7 DA EX, RT PAC-96BO-064-01 B 2b 6 UNIM UNI F 1 1–2 PO EX

PAC-96BO-064-02 B 2b 6 UNIM UNI F 1 2–3 PO EX, RT

PAC-96BO-064-03 B 2b 6 ODO MTP DE L 1 U 1.9 1.7 DA EX PAC-96BO-064-04 B 2b 6 ODO HUM DS R 1 5.1 2.1 DA EX, CRK

PAC-96BO-066-01 B 2+2b 5 UNIM UNI F 1 1–2 PO EX

PAC-96BO-066-02 B 2+2b 6 ODO HUM DS R 1 4.3 1.9 PO EX, RT

PAC-96BO-067-01 B 12 7 UNI UNI F 4 <1 IT EX, RT

PAC-96BO-067-02 B 12 7 UNI UNI F 1 1–2 PO EX

PAC-96BO-067-03 B 12 7 ODO RIB POST L 1 3.3 0.8 DA EX, STG

PAC-96BO-067-04 B 12 7 ARM TIB DE+MS+DS R 1 FU 4.2 1.7 PO EX PAC-96BO-067-05 B 12 7 AVES LBF F 1 2.3 0.6 DA RT

PAC-96BO-068-01 B 11 7 UNIM UNI F 1 1–2 PO EX

PAC-96BO-068-02 B 11 7 UNIM UNI F 1 1–2 SD EX

PAC-96BO-068-03 B 11 7 UNIM UNI F 1 1–2 SD EX

PAC-96BO-068-04 B 11 7 MTM-LTM UNI F 1 2.5 1.2 PO EX, RT

212

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-96BO-068-05 B 11 7 MTM-LTM UNI F 1 2.2 1.3 PO EX, RT

PAC-96BO-068-06 B 11 7 MTM-LTM LBF F 1 2.1 1.6 PO EX, RT

PAC-96BO-084-01 B 8 6B UNI UNI F 3 <1 PO EX

PAC-96BO-084-02 B 8 6B UNIM UNI F 3 1–2 PO EX

PAC-96BO-084-03 B 8 6B UNIM UNI F 1 2–3 PO EX, RT

PAC-96BO-084-04 B 8 6b MTM-LTM LBF F 1 3.5 1.1 PO EX, RT

PAC-96BO-084-05 B 8 6B OCE HUM DE R 1 FU 1.4 1.2 PO EX, STG PAC-96BO-084-05 B 8 6b MTM-LTM LBF F 1 3.9 0.9 PO EX, RT

PAC-96BO-084-06 B 8 6B ODO MTT S 1 3.4 1.4 PO EX, RT

PAC-96BO-099-01 B 1 8 UNI UNI F 4 <1 SD EX

PAC-96BO-099-02 B 1 8 UNIM UNI F 4 1–2 DA EX

PAC-96BO-099-03 B 1 8 UNIM UNI F 2 1–2 PO EX, RT

PAC-96BO-099-05 B 1 8 UNIM UNI F 1 2–3 PO EX

PAC-96BO-099-06 B 1 8 MTM-LTM LBF F 1 3.1 1.4 DA EX, RT

PAC-96BO-099-07 B 1 8 MTM-LTM LBF F 1 5.5 0.8 DA EX, RT

PAC-96BO-099-08 B 1 8 ODO MTC PE L 1 FU 1.5 1.4 DA EX PAC-96BO-099-09 B 1 8 ODO MTP S 1 3.6 0.7 DA EX, RT

PAC-96BO-099-10 B 1 8 ODO PHA DE+DS 1 FU 1.4 1.2 SD EX

PAC-96BO-099-11 B 1 8 RAB ULN PE R 1 FU 1.4 0.7 SD RT, STG PAC-96BO-099-12 B 1 8 TES SHL F 1 1.0 0.5 SD EX, CRK

PAC-96BO-099-13 B 1 8 TES SHL F 1 2.4 1.2 SD EX, RT, STG, CT

PAC-96BO-099-14 B 1 8 MTM LBF F 1 1.6 1.4 DA EX, RT, GN, BUR

PAC-96BO-100-01 B 8 8 UNIM UNI F 1 2–3 SD EX, STG

PAC-96BO-100-02 B 8 8 UNIM UNI F 1 1–2 PO EX, STG

PAC-96BO-103-01 B 6 7 UNI UNI F 6 <1 DA EX

PAC-96BO-103-02 B 6 7 UNIM UNI F 3 1–2 DA EX

PAC-96BO-103-03 B 6 7 UNIM UNI F 4 2–3 DA EX

213

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-96BO-103-04 B 6 7 UNIM UNI F 4 1–2 DA EX

PAC-96BO-103-05 B 6 7 UNIM UNI F 1 1–2 PO EX

PAC-96BO-103-09 B 6 7 LTM UNI F 1 2.5 2.2 PO EX, RT

PAC-96BO-103-10 B 6 7 LTM LBF F 1 2.4 1.6 PO EX, RT, CT

PAC-96BO-103-11 B 6 7 ODO TRQ W L 1 FU 2.2 1.6 SD EX, STG PAC-96BO-103-12 B 6 7 ARM ULN PE+PS+MS R 1 FU 2.6 1.1 DA EX, RT, STG PAC-96BO-103-13 B 6 7 ARM CDV F AX 1 U 1.6 0.6 DA EX, RT PAC-96BO-103-14 B 6 7 ARM TIB DS+MS R 1 3.4 1.2 DA EX, RT

PAC-96BO-103-15 B 6 7 ARM FIB DS+MS R 1 2.4 1.6 DA EX, RT

PAC-96BO-105-01 B 8 7 UNI UNI F 1 <1 PO EX

PAC-96BO-105-02 B 8 7 UNIM UNI F 2 1–2 DA EX

PAC-96BO-105-03 B 8 7 FISH VER W-S AX 1 1.3 0.5 PO EX

PAC-96BO-107-01 B 8 6B UNI UNI F 1 1–2 SD RT, BUR

PAC-96BO-108-01 B 11 6B UNIM UNI F 1 1–2 PO EX

PAC-96BO-108-02 B 11 6B UNIM UNI F 1 2–3 PO EX, RT

PAC-96BO-108-03 B 11 6B UNIM UNI F 1 4–5 PO EX, RT

PAC-96BO-108-04 B 11 6b STM-MTM LBF F 1 2.2 0.5 PO EX, RT, CRK, STG

PAC-96BO-108-05 B 11 6b LTM LBF F 1 3.0 1.5 PO EX, RT

PAC-96BO-108-06 B 11 6B ODO NVB F L 1 FU 2.9 2.5 PO EX, RT, STG PAC-96BO-109-01 B 11 6B UNIM UNI F 1 2–3 PO EX, RT

PAC-96BO-109-02 B 11 6b LTM LBF F 1 3.5 1.6 PO EX, RT

PAC-96BO-109-02 B 11 6B UNIM UNI F 1 <1 SD CRK

PAC-96BO-109-03 B 11 6b STM-MTM LBF F 1 2.0 0.7 PO EX

PAC-96BO-110-01 B 6 6 UNIM UNI F 3 <1 PO EX

PAC-96BO-110-02 B 6 6 UNIM UNI F 1 1–2 PO EX

PAC-96BO-110-03 B 6 6 UNIM UNI F 1 2–3 PO EX

PAC-96BO-110-04 B 6 6 UNIM UNI F 1 4–5 DA EX

214

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-96BO-110-05 B 6 6 ODO ULN PE R 1 U 2.4 1.3 PO EX PAC-96BO-110-06 B 6 6 LTM LBF F 1 4.6 1.8 PO EX, RT

PAC-96BO-110-07 B 6 6 TES SHL Marginal 1 1.2 1.1 PO EX

PAC-96BO-111-01 B 8 7 UNI UNI F 4 <1 PO EX

PAC-96BO-112-01 B 8 7 UNI UNI F 2 <1 DA EX

PAC-96BO-112-02 B 8 7 UNIM UNI F 1 1–2 SD EX

PAC-96BO-112-03 B 8 7 ARM FEM MS L 1 2.5 1.4 PO EX, RT

PAC-96BO-113-01 B 10 7 UNIM UNI F 1 1–2 DA EX

PAC-96BO-113-02 B 10 7 PAC ULN PE+PS R 1 FU 3.5 1.2 SD EX, RT PAC-96BO-113-03 B 10 7 UNI UNI F 1 <1 PO EX, STG

PAC-96BO-116-01 B 3 6 UNI UNI F 7 <1 PO EX

PAC-96BO-116-02 B 3 6 UNI UNI F 7 1–2 PO EX

PAC-96BO-117-01 B 6 6 UNIM UNI F 2 1–2 PO EX

PAC-96BO-117-02 B 6 6 LTM LBF F 1 3.1 1.5 PO EX, RT

PAC-96BO-117-03 B 6 6 COL VER W AX 1 FU 0.9 0.8 SD

PAC-96BO-118-01 B 3 6 UNI UNI F 1 1–2 PO EX

PAC-96BO-118-02 B 3 6 STM LBF F 1 1.1 0.5 PO SH

PAC-96BO-118-03 B 3 6 MTM LBF F 1 0.9 0.9 DA EX, RT

PAC-96BO-119-01 B 4 6 UNIM UNI F 1 2–3 DA EX

PAC-96BO-119-02 B 4 6 LTM LBF F 1 4.0 1.4 PO EX, RT

PAC-96BO-120-01 B 2c 6 UNIM UNI F 1 1–2 DA EX

PAC-96BO-121-01 B 9 6 UNI UNI F 4 <1 DA EX

PAC-96BO-121-02 B 9 6 UNIM UNI F 1 1–2 DA EX, RT

PAC-96BO-121-03 B 9 6 LTM LBF F 1 3.2 1.6 SD EX, STG

PAC-96BO-121-04 B 9 6 UNIM UNI F 5 1–2 PO EX

PAC-96BO-121-05 B 9 6 UNI UNI F 2 2–3 DA EX

PAC-96BO-132-01 B 6 8 MTM-LTM LBF F 1 2.6 1.4 PO EX, RT

215

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-96BO-142-01 B 1 6 UNI UNI F 2 <1 DA EX

PAC-96BO-142-02 B 1 6 UNIM UNI F 1 1–2 DA EX

PAC-96BO-142-03 B 1 6 TES SHL F 3 0.8 0.7 PO EX

PAC-96BO-142-04 B 1 6 AVES STX F AX 1 1.0 0.5 PO EX

PAC-96BO-142-05 B 1 6 GOP TIB PE+PS L 1 U 0.8 0.7 PO EX PAC-96BO-143-01 B 2c 6 UNIM UNI F 1 3–4 PO EX

PAC-96BO-143-02 B 2c 6 TAY PM3 UP L 1 1.0 0.9 DA EX

PAC-96BO-148-01 B 5 7 ARM TIB DS+MS R 1 2.5 1.1 PO EX, RT, STG

PAC-96BO-149-01 B 5 7 UNIM UNI F 1 1–2 PO EX, RT

PAC-96BO-151-01 B 6 7 UNIM UNI F 2 2–3 DA EX

PAC-96BO-158-01 B 8 6 UNI UNI F 2 <1 DA EX

PAC-96BO-158-02 B 8 6 CAN CN UP L 1 1.5 0.5 DA EX, RT

PAC-96BO-160-01 B 10 6 TAY PH2 W 1 FU 1.8 1.0 SD EX, RT

PAC-96BO-161-01 B 10 7 STM-MTM LBF F 1 1.5 0.7 DA EX, RT, CRK, GN

PAC-96BO-162-01 B 11 6B ODO ILM F L 1 3.8 1.5 PO EX, STG

PAC-96BO-163-01 B 11 7 UNI UNI F 5 <1 PO EX

PAC-96BO-163-02 B 11 7 UNIM UNI F 2 2–3 PO EX, RT

PAC-96BO-163-03 B 11 7 UNIM UNI F 2 2–3 PO EX, RT

PAC-96BO-163-04 B 11 7 UNIM UNI F 4 1–2 DA EX

PAC-96BO-163-05 B 11 7 UNIM UNI F 4 2–3 DA EX

PAC-96BO-163-06 B 11 7 UNIM UNI F 2 2–3 SD EX

PAC-96BO-163-08 B 11 7 STM LBF F 1 1.5 0.4 DA EX, STG

PAC-96BO-163-09 B 11 7 LTM LBF F 1 3.6 0.9 SD RT

PAC-96BO-163-10 B 11 7 LTM LBF F 1 3.7 1.4 DA EX, RT

PAC-96BO-163-11 B 11 7 LTM LBF F 1 3.9 1.4 PO EX, RT

PAC-96BO-163-12 B 11 7 LTM LBF F 1 5.5 1.6 SD EX, RT

PAC-96BO-163-13 B 11 7 ARM SHL F 1 1.4 0.5 SD EX

216

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-96BO-163-14 B 11 7 MAZ MAL W L 1 FU 1.1 0.9 DA EX, RT PAC-96BO-163-15 B 11 7 ODO TIB MS R 1 4.7 1.6 DA EX, RT, STG

PAC-96BO-163-16 B 11 7 ODO PH3 PE+PS 1 1.8 1.1 PO EX

PAC-96BO-163-17 B 11 7 ICT DRS ANT AX 1 1.7 0.5 SD RT, STG

PAC-96BO-163-18 B 11 7 FISH DRS ANT AX 1 1.4 0.5 DA EX, RT

PAC-96BO-163-20 B 11 7 ODO SES W 1 FU 1.2 1.0 DA EX

PAC-96BO-164-01 B 11 7B UNIM UNI F 2 1–2 DA EX

PAC-96BO-164-02 B 11 7B TES SHL F 1 1.2 0.8 DA EX, RT, GN

PAC-97BO-007-01 B 97-SU1 6 UNIM UNI F 1 <1 PO EX

PAC-97BO-007-02 B 97-SU1 6 UNIM UNI F 4 1–2 PO EX

PAC-97BO-007-03 B 97-SU1 6 UNIM UNI F 1 1–2 PO SH, CRK

PAC-97BO-011-01 D 97D1 7 UNIM LBF F 1 4.1 1.9 DA EX, RT

PAC-97BO-011-02 D 97D1 7 UNIM UNI F 1 2.2 1.2 SD RT, BUR

PAC-97BO-016-01 B 97-U1 6 UNI UNI F 1 <1 PO EX

PAC-97BO-016-02 B 97-U1 6 SER CDV W-S AX 1 1.0 0.7 DA EX

PAC-97BO-016-03 B 97-U1 6 TAY PH1 PE+PS 1 FU 0.7 0.5 PO EX

PAC-97BO-021-01 B 97-SU4 6 MTM UNI F 1 1.3 0.4 PO BUR

PAC-97BO-025-01 B 97-RU1 6C UNIM UNI F 1 <1 SD EX

PAC-97BO-027-01 B 97-SU3 6 UNIM LBF F 1 1.8 0.6 SD EX

PAC-97BO-029-01 B 97-U1 6 UNI UNI F 1 <1 PO EX

PAC-97BO-029-02 B 97-U1 6 UNIM UNI F 1 1–2 PO EX

PAC-97BO-029-03 B 97-U1 6 ODO RIB F R 1 2.0 0.7 DA EX, CT

PAC-97BO-029-04 B 97-U1 6 ODO MO LO L 1 1.2 0.5 PO EX

PAC-97BO-031-01 B 97-U3 6 UNIM UNI F 1 <1 PO EX

PAC-97BO-031-02 B 97-U3 6 UNIM UNI F 3 1–2 PO EX

PAC-97BO-031-03 B 97-U3 6 MAZ LUN W L 1 FU 1.3 1.0 PO EX PAC-97BO-052-01 B 97-U1 8 UNI UNI F 3 <1 DA EX

217

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-97BO-052-02 B 97-U1 8 UNIM UNI F 1 2–3 DA EX

PAC-97BO-052-03 B 97-U1 8 ODO CAL DE+MS R 1 FU 4.3 1.6 DA EX

218

Table A.2 Pacbitun dataset for the late Middle Preclassic period

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state P95-00-02 B 1 5 UNIM UNI F 2 1–2 SD EX, CRK

P95-00-03 B 1 5 UNIM UNI F 3 2–3 DA EX

P95-00-04 B 1 5 UNIM UNI F 1 3–4 PO EX, RT

P95-00-05 B 1 5 UNIM UNI F 1 1.1 0.7 IT BUR

P95-00-06 B 1 5 LTM LBF F 1 2.9 1.4 PO EX, RT

P95-00-07 B 1 5 LTM LBF F 1 3.7 1.0 PO EX, RT

P95-00-08 B 1 5 LTM LBF F 1 5.6 1.2 PO EX, RT, STG, PN

P95-02-01 B 1 4 LTM LBF F 1 6.3 1.9 PO EX

P95-03-01 B 1 3 UNIM UNI F 1 1.2 PO EX

P95-04-01 B 1 5 UNIM UNI F 1 1.2 PO EX

P95-05-01 B 1 5 UNIM UNI F 1 2.2 PO EX

P95-06-01 B 1 4 UNI UNI F 4 <1 PO EX

P95-06-02 B 1 4 UNI UNI F 1 1–2 PO EX

P95-06-03 B 1 4 UNI UNI F 2 2–3 PO EX

P95-06-04 B 1 4 LTM LBF F 1 6.0 2.1 DA EX, RT, SH, STG

P95-06-05 B 1 4 IGU VER F AX 1 1.0 0.9 DA EX, RT, STG

P95-07-01 B 1 3 ODO CEV CAU AX 1 2.6 2.2 DA EX, RT

P95-07-02 B 1 3 UNIM UNI F 1 1.0 PO EX

P95-07-03 B 1 3 LTM LBF F 1 3.8 1.0 PO EX, RT, STG

P95-08-01 B 1 5 UNI UNI F 1 1–2 PO EX

P95-09-01 B 1 3 UNIM UNI F 5 <1 PO EX

P95-09-02 B 1 3 UNIM UNI F 13 1–2 PO EX, STG

P95-09-03 B 1 3 UNIM LBF F 3 1–2 PO EX

P95-09-04 B 1 3 UNIM LBF F 4 2–3 PO EX

P95-09-05 B 1 3 ODO AST PRO L 1 FU 3.0 2.2 PO EX, STG, CRK

219

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state P95-09-06 B 1 3 ODO HUM DE R 1 FU 3.8 1.5 PO EX, CRK P95-09-07 B 1 3 MAR NAV W R 1 FU 1.0 0.8 PO EX P95-09-08 B 1 3 COL VER W AX 1 FU 1.0 1.0 PO EX P95-09-09 B 1 3 COL VER W AX 1 FU 1.0 1.0 PO EX P95-09-10 B 1 3 LTM UNI F 1 3.6 1.2 PO EX

P95-09-11 B 1 3 LTM LBF F 1 2.8 1.4 PO EX, RT

P95-09-12 B 1 3 MTM LBF F 1 2.6 0.9 PO STG P95-09-14 TBT DS R 1 2.4 0.7 B 1 3 AVES SD RT, STG P95-09-14 B 1 3 LTM LBF F 1 4.7 1.3 PO EX, RT, CRK

P95-11-01 B 1 4 UNIM UNI F 6 <1 PO EX

P95-11-02 B 1 4 ODO NVB W R 1 FU 2.7 2.4 DA RX, RT, STG P95-11-03 B 1 4 ODO HUM DE+DS R 1 5.7 2.3 DA EX

P95-11-04 B 1 4 ODO GC F R 1 FU 1.7 0.8 SD EX, STG P95-12-01 B 1 3 UNIM UNI F 1 2.0 DA EX

P95-12-02 B 1 3 ARM SHL F 1 0.8 0.6 IT

P95-13-01 B 1 3 UNIM UNI F 1 1.6 PO EX

P95-14-01 B 1 ext. 3 UNIM UNI F 1 1.4 PO EX

P95-15-01 B 1 3 UNIM UNI F 1 1–2 PO EX

P95-15-02 B 1 3 ARM CAL F L 1 FU 1.9 0.8 DA EX P95-16-01 B 1 3 UNI UNI F 4 <1 PO EX

P95-16-02 B 1 3 UNIM UNI F 1 1.7 PO EX

P95-16-04 B 1 3 LTM LBF F 1 7.9 1.9 PO EX, STG

P95-17-01 B 1 5 UNIM UNI F 2 1–2 DA EX

P95-17-02 B 1 5 UNIM LBF F 1 1.7 DA EX, BUR

P95-18-01 B 1 3 UNI UNI F 3 1–2 PO EX

P95-18-02 B 1 3 UNI UNI F 1 2.8 1.3 PO EX

P95-18-03 B 1 3 LTM UNI F 1 1.9 1.6 PO EX

220

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state P95-18-04 B 1 3 ODO TIB PE L 1 2.2 1.4 DA EX

P95-18-05 B 1 3 ODO ILM F R 1 3.7 1.6 PO EX

P95-19-01 B UNIM UNI F 1 <1 PO EX

P95-19-02 B UNIM UNI F 1 1–2 PO EX, CRK

P95-19-03 B UNIM UNI F 1 2–3 PO EX

P95-19-04 B UNIM UNI F 2 3–4 PO EX

P95-19-05 B UNIM LBF F 1 3–4 PO EX

P95-19-06 B UNIM UNI F 1 4–5 PO EX

P95-20-01 B 1 3 UNIM UNI F 2 1–2 PO EX

P95-20-02 B 1 3 UNIM UNI F 1 2–3 PO EX

P95-20-04 B 1 3 UNI UNI F 2 1–2 PO EX

P95-21-01 B 1 3 UNIM UNI F 9 1–2 PO EX, STG

P95-21-02 B 1 3 UNIM LBF F 2 2–3 PO EX, RT

P95-21-03 B 1 3 UNIM UNI F 1 1–2 PO EX, GN

P95-21-04 B 1 3 ARM FEM PE L 1 FU 1.0 0.9 PO EX, GN P95-21-05 B 1 3 TAY PH1 W 1 FU 1.3 0.6 PO EX

P95-21-06 B 1 3 ODO CAL DE+DS R 1 FU 3.4 1.9 PO EX P95-21-07 B 1 3 MTM-LTM LBF F 1 3.3 0.9 PO EX, RT, BUR

P95-21-09 B 1 3 ODO SES W 1 FU 0.9 0.9 PO EX

P95-22-01 B 1 3 UNIM UNI F 7 <1 PO EX

P95-22-02 B 1 3 UNIM UNI F 10 1–2 PO EX

P95-22-03 B 1 3 STM LBF F 1 2.0 0.4 PO EX, RT

P95-22-04 B 1 3 STM LBF F 1 1.5 0.4 PO EX, RT

P95-22-05 B 1 3 DOG MO1 LO L 1 2.2 2.2 PO EX, CRK, STG

P95-22-06 B 1 3 MAZ MAL W L 1 FU 0.9 0.8 PO EX, RT P95-22-07 B 1 3 ODO MO3 LO R 1 2.2 0.9 DA EX

P95-22-08 B 1 3 MTM-LTM LBF F 1 2.0 0.9 PO EX

221

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state P95-23-01 B 1 3 UNIM UNI F 6 <1 PO EX

P95-23-02 B 1 3 UNIM UNI F 2 1–2 PO EX

P95-23-03 B 1 3 UNIM UNI F 2 2–3 PO EX

P95-23-04 B 1 3 UNIM UNI F 1 1.9 0.9 SD BUR

P95-23-05 B 1 3 CER TRV F AX 1 1.1 1.0 PO EX

P95-23-06 B 1 3 LTM VER F AX 1 2.0 1.7 DA EX

P95-24-01 B 1 3 UNIM UNI F 2 1–2 PO EX

P95-24-02 B 1 3 UNIM UNI F 3 2–3 PO EX

P95-24-03 B 1 3 MTM LBF F 1 2.6 0.9 PO EX, PN

P95-24-04 B 1 3 MTM LBF F 1 2.0 1.1 PO EX, RT

P95-25-01 B 1 4 UNIM UNI F 16 <1 PO EX

P95-25-02 B 1 4 UNIM UNI F 7 1–2 PO EX

P95-25-03 B 1 4 UNIM UNI F 3 1–2 PO EX

P95-25-04 B 1 4 MTM-LTM LBF F 1 3.4 1.0 PO EX, BUR

P95-25-06 B 1 4 TAY RUL PE+PS R 1 FU 5.2 2.0 DA EX, RT, CRK P95-25-07 B 1 4 MAZ PH1 DE+DS+MS 1 FU 2.5 0.9 PO EX, RT

P95-25-08 B 1 4 ARM CAL PE+PS R 1 FU 1.4 1.1 DA EX P95-26-01 B UNIM UNI F 6 <1 PO EX

P95-26-02 B UNIM UNI F 6 1–2 PO EX, STG

P95-26-03 B UNIM UNI F 7 1–2 PO EX

P95-26-04 B UNIM UNI F 2 2–3 PO EX

P95-26-05 B UNIM UNI F 3 2–3 SD RT, BUR

P95-26-06 B ARM SHL F 1 0.9 0.5 DA EX, STG

P95-26-06 B ODO FEM PS L 1 4.7 1.3 PO EX

P95-26-07 B TES SHL F 1 2.1 1.5 PO EX

P95-27-01 B UNIM UNI F 1 2–3 PO EX

P95-27-02 B UNIM UNI F 1 3–4 PO EX

222

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state P95-28-01 B 1 4 UNIM UNI F 1 1–2 DA EX

P95-29-01 B 1 3+4 UNIM UNI F 1 2–3 SD STG

P95-29-02 B 1 3+4 UNIM UNI F 1 1–2 PO EX

P95-30-01 B BUF HUM W L 1 FU 1.8 0.5 IT

P95-30-03 B ODO DP4 LO R 1 1.1 1.1 SD EX

P95-30-04 B UNIM UNI F 1 1–2 DA EX, RT, BUR

P95-30-05 B TES HUM F L 1 FU 1.5 0.6 IT RT, STG

P95-32-01 B UNIM UNI F 3 1–2 SD

P95-32-02 B MAZ TIB PS+MS L 1 8.8 2.9 SD EX, RT, STG, GN

P95-32-03 B ARM RAD PE+PS+MS R 1 FU 1.8 0.5 SD EX

P95-33-01 B 1 4 UNIM UNI F 8 <1 PO EX

P95-33-02 B 1 4 UNIM UNI F 3 1–2 PO EX

P95-33-03 B 1 4 UNIM UNI F 4 1–2 PO EX

P95-33-04 B 1 4 MTM-LTM SKL F 1 2.4 1.5 PO EX, RT

P95-33-05 B 1 4 UNIM LBF F 1 1.2 0.7 DA EX, BUR

P95-33-06 B 1 4 UNIM UNI F 1 4–5 PO EX, STG

P95-33-07 B 1 4 ODO PH3 W 1 FU 3.0 1.3 SD EX

P95-33-09 B 1 4 TAP PM3 LO L 1 1.8 1.5 PO EX, STG

P95-33-10 B 1 4 GOP IN1 UP R 1 1.8 1.4 SD EX, RT

P95-33-11 B 1 4 MTM PH3 W 1 FU 1.6 0.7 DA EX

P95-33-12 B 1 4 IGU ULN DE+DS+MS R 1 FU 1.8 0.3 SD EX P95-34-01 B UNIM UNI F 1 1–2 PO EX

P95-34-02 B MAZ PH1 DE+DS+MS 1 FU 1.9 0.9 PO EX, GN

P95-41-01 B 1 4 UNI UNI F 1 1–2 PO EX

P95-75-01 B 1 3 UNIM LBF F 1 1.1 0.6 SD BUR

P95-90-01 B CER MTT PS R 1 3.3 0.8 PO EX, STG

PAC-08BO-001-01 B 1 4 UNIM LBF F 1 2.1 1.9 PO EX

223

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-08BO-002-01 B 1 4 UNI UNI F 1 1–2 PO EX

PAC-08BO-003-01 B 1 4 ARM TIB DS R 1 2.2 0.8 PO EX, RT, STG

PAC-08BO-004-01 B 1 4 UNIM UNI F 1 1–2 PO EX

PAC-08BO-005-01 B 1 4 UNIM UNI F 3 <1 PO EX

PAC-08BO-005-02 B 1 4 UNIM UNI F 1 1–2 SD EX

PAC-08BO-006-01 B 1 5 UNIM UNI F 1 <1 PO EX

PAC-08BO-006-02 B 1 5 UNIM UNI F 2 1–2 PO EX

PAC-08BO-006-03 B 1 5 ODO TRV F L 1 1.6 1.4 PO EX, STG, CRK

PAC-08BO-006-04 B 1 5 LTM VER F AX 1 U 1.7 1.6 PO EX, RT PAC-08BO-007-01 B 1 5 UNIM UNI F 1 <1 PO EX

PAC-08BO-007-02 B 1 5 UNIM UNI F 1 2–3 DA EX

PAC-08BO-008-01 B 1 5 UNIM UNI F 5 <1 PO EX

PAC-08BO-009-01 B 1 5 UNIM UNI F 2 <1 PO EX

PAC-08BO-009-02 B 1 5 UNIM UNI F 1 1–2 PO EX

PAC-08BO-009-03 B 1 5 VIP VER F AX 1 FU 0.7 0.3 PO EX PAC-08BO-010-02 B 1 5 UNIM UNI F 2 1–2 DA EX

PAC-08BO-010-03 B 1 5 FISH VER W-S AX 1 0.6 0.5 SD EX

PAC-08BO-011-01 B 1 5 UNIM UNI F 1 4–5 SD EX, RT, STG

PAC-08BO-011-02 B 1 5 ODO MTP S 1 3.6 1.2 DA SH, CRK

PAC-08BO-012-01 B 1 5 UNIM UNI F 1 1–2 PO EX

PAC-08BO-012-02 B 1 5 UNIM UNI F 1 1–2 DA EX, STG

PAC-08BO-013-01 B 1 5 UNIM UNI F 1 1–2 PO EX

PAC-08BO-013-02 B 1 5 UNIM UNI F 1 1–2 PO EX

PAC-08BO-013-03 B 1 5 STM LBF F 1 1.9 0.5 PO EX, RT

PAC-08BO-016-01 B 2 4 UNIM UNI F 10 <1 PO EX

PAC-08BO-016-02 B 2 4 UNIM UNI F 4 1–2 PO EX

PAC-08BO-016-03 B 2 4 VIP VER F AX 1 FU 1.8 0.8 PO EX

224

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-08BO-017-01 B 2 4 UNIM UNI F 1 2–3 SD STG

PAC-08BO-018-01 B 2 5 UNIM UNI F 1 <1 PO EX

PAC-08BO-018-02 B 2 5 UNIM UNI F 2 1–2 PO EX

PAC-08BO-018-03 B 2 5 UNIM UNI F 2 2–3 PO EX, RT

PAC-08BO-019-01 B 2 5 UNIM UNI F 3 <1 PO EX

PAC-08BO-019-02 B 2 5 UNIM UNI F 2 1–2 PO EX

PAC-08BO-021-01 B 1 4 UNIM UNI F 1 2–3 SD EX

PAC-08BO-023-01 B 2 5 UNI UNI F 1 1–2 SD

PAC-08BO-023-02 B 2 5 AVES LBF F 1 2.1 0.5 DA EX, RT

PAC-09BO-001-01 B 1 4 UNIM UNI F 6 <1 PO EX

PAC-09BO-001-02 B 1 4 UNIM UNI F 2 1–2 PO EX

PAC-09BO-001-03 B 1 4 UNIM UNI F 2 2–3 PO EX, RT

PAC-09BO-001-04 B 1 4 UNIM UNI F 1 1–2 PO EX, CRK

PAC-09BO-002-01 B 1 5 UNI UNI F 1 <1 PO EX

PAC-09BO-002-02 B 1 5 UNIM LBF F 1 1.7 0.6 PO EX, RT

PAC-09BO-002-03 B 1 5 UNIM UNI F 1 2–3 PO EX, RT

PAC-09BO-002-04 B 1 5 ODO NVB F L 1 FU 1.4 1.0 PO EX PAC-09BO-002-05 B 1 5 STM LBF F 1 1.5 0.4 PO EX, RT

PAC-09BO-002-06 B 1 5 CER MTP S 1 3.8 1.3 PO EX, STG, PN

PAC-09BO-003-01 B 1 5 UNI UNI F 4 <1 PO EX

PAC-09BO-003-02 B 1 5 UNIM UNI F 10 1–2 PO EX

PAC-09BO-003-03 B 1 5 UNIM UNI F 3 2–3 PO EX

PAC-09BO-003-06 B 1 5 UNIM LBF F 1 1.6 1.3 DA EX, RT, BUR

PAC-09BO-003-07 B 1 5 ODO PH1 DE+DS+MS+PS 1 FU 3.6 1.3 PO EX

PAC-09BO-003-08 B 1 5 ODO PH1 DE 1 FU 1.2 1.0 PO EX, CRK

PAC-09BO-003-09 B 1 5 ODO PH2 DE 1 FU 1.8 1.4 SD EX, BUR

PAC-09BO-003-10 B 1 5 ODO PH2 PE+PS 1 FU 1.9 1.4 SD EX, BUR

225

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-09BO-003-11 B 1 5 PUM HUM DE L 1 FU 5.2 4.9 PO EX, RT PAC-09BO-003-12 B 1 5 ODO HUM DS R 1 2.8 1.6 DA EX, STG

PAC-09BO-003-13 B 1 5 GOP IN1 LO 1 1.8 0.4 SD EX, RT

PAC-09BO-003-14 B 1 5 STM LBF F 1 1.1 0.4 PO EX, RT, STG

PAC-09BO-003-15 B 1 5 MTM-LTM LBF F 1 4.7 1.1 DA EX, RT

PAC-09BO-004-01 B 1 5 UNI UNI F 1 1–2 PO EX, STG, CRK

PAC-09BO-005-01 B 1 5 UNI UNI F 19 <1 PO EX

PAC-09BO-005-02 B 1 5 UNIM UNI F 7 1–2 PO EX

PAC-09BO-005-03 B 1 5 UNIM UNI F 1 3–4 PO EX, RT

PAC-09BO-005-04 B 1 5 UNIM UNI F 1 3–4 DA EX, RT

PAC-09BO-005-05 B 1 5 CER TRV CEN AX 1 FU 1.6 1.0 PO EX PAC-09BO-005-06 B 1 5 LTM LMV F AX 1 FU 3.3 2.1 DA EX PAC-09BO-005-07 B 1 5 LTM LMV F AX 1 FU 2.2 1.2 DA EX PAC-09BO-005-08 B 1 5 MTM-LTM VER CEN AX 1 FU 1.1 0.7 SD RT PAC-09BO-005-11 B 1 5 LTM VER CEN AX 1 FU 1.8 1.1 DA EX, GN PAC-09BO-006-01 B 1 5 UNI UNI F 1 <1 DA EX

PAC-09BO-006-02 B 1 5 STM LBF F 1 1.2 0.3 PO EX

PAC-09BO-011-01 B 2 4 UNIM UNI F 1 1–2 PO EX, RT

PAC-09BO-012-01 B 2 5 UNIM UNI F 1 <1 PO EX, RT

PAC-09BO-013-01 B 2 5 UNIM UNI F 1 <1 PO EX

PAC-09BO-013-02 B 2 5 ODO MTP DE 1 U 2.0 1.8 DA EX, GN

PAC-09BO-014-01 B 2 5 UNI UNI F 6 <1 PO EX

PAC-09BO-014-02 B 2 5 UNIM UNI F 3 1–2 PO EX

PAC-09BO-014-03 B 2 5 UNI UNI F 1 1–2 DA EX

PAC-09BO-014-04 B 2 5 UNIM LBF F 1 2.2 1.2 DA EX, RT

PAC-09BO-014-05 B 2 5 ODO PUB ACE L 1 FU 2.2 1.9 PO EX PAC-09BO-014-06 B 2 5 LTM LBF F 1 5.0 1.6 PO EX, CRK, STG

226

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-09BO-015-01 B 2 5 UNIM UNI F 3 1–2 PO EX

PAC-09BO-015-03 B 2 5 ODO SES W 1 FU 1.0 1.0 SD EX

PAC-09BO-016-01 B 2 5 UNI UNI F 1 <1 PO EX

PAC-09BO-016-02 B 2 5 UNIM UNI F 7 1–2 PO EX

PAC-09BO-016-04 B 2 5 UNIM UNI F 4 1–2 DA EX, RT

PAC-09BO-016-05 B 2 5 UNIM LBF F 1 2–3 DA EX, RT

PAC-09BO-016-06 B 2 5 UNIM UNI F 2 2–3 DA EX, STG

PAC-09BO-016-07 B 2 5 ODO RAD PS R 1 2.9 1.0 DA EX, STG

PAC-09BO-016-08 B 2 5 ODO CAP W R 1 FU 1.7 1.3 PO EX, GN PAC-09BO-016-09 B 2 5 MAZ PH3 W 1 FU 1.8 1.0 DA EX

PAC-09BO-016-10 B 2 5 ODO MTP DE 1 FU 1.2 1.2 PO EX

PAC-09BO-016-11 B 2 5 ODO PH3 W 1 FU 2.4 1.5 PO EX

PAC-09BO-016-12 B 2 5 AGO MTT 4 PE+PS+MS R 1 FU 2.0 0.5 PO EX PAC-09BO-016-13 B 2 5 TES SHL F 1 1.1 1.1 PO SH

PAC-09BO-016-14 B 2 5 LTM UNI F 1 3.9 0.9 DA EX, RT

PAC-09BO-016-15 B 2 5 LTM UNI F 1 2.6 1.8 PO EX

PAC-09BO-017-01 B 2 5 UNIM LBF F 1 3.2 1.4 PO EX, RT

PAC-09BO-022-01 B 3 4 UNIM UNI F 5 1–2 PO EX, RT

PAC-09BO-022-02 B 3 4 UNIM LBF F 1 1.4 1.2 PO EX, RT, BUR

PAC-09BO-023-01 B 3 5 UNI UNI F 1 <1 PO EX, RT

PAC-09BO-024-01 B 3 5 UNI UNI F 10 <1 PO EX

PAC-09BO-024-02 B 3 5 UNIM UNI F 22 1–2 PO EX

PAC-09BO-024-03 B 3 5 UNIM UNI F 7 2–3 PO EX, RT

PAC-09BO-024-06 B 3 5 UNIM UNI F 5 1–2 PO EX

PAC-09BO-024-07 B 3 5 ODO PH1 DE+DS+MS+PS 1 FU 3.5 1.1 PO EX, GN

PAC-09BO-024-08 B 3 5 ODO HUM DE L 1 FU 2.9 2.7 PO EX PAC-09BO-024-09 B 3 5 ODO PH1 PE 1 1.8 0.7 DA EX

227

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-09BO-024-10 B 3 5 ODO PM3-4 UP R 1 1.8 0.9 PO EX

PAC-09BO-024-11 B 3 5 LTM LBF F 1 4.6 1.3 PO EX

PAC-09BO-024-12 B 3 5 LTM UNI F 1 2.7 1.7 PO EX

PAC-09BO-024-13 B 3 5 LTM UNI F 1 3.2 1.9 PO EX

PAC-09BO-025-01 B 1 4 UNIM UNI F 4 1–2 PO EX, RT

PAC-09BO-026-01 B 1 4 UNI UNI F 6 <1 PO EX

PAC-09BO-026-02 B 1 4 UNIM UNI F 1 1–2 PO EX, RT

PAC-09BO-026-03 B 1 4 UNIM UNI F 1 1–2 PO EX, RT

PAC-09BO-027-01 B 1 4 UNIM UNI F 1 2–3 DA EX, RT

PAC-09BO-028-01 B 1 4 UNI UNI F 1 1–2 PO EX

PAC-09BO-029-01 B 3 5 UNIM UNI F 1 1–2 PO EX, RT

PAC-09BO-033-01 B 4 4 UNIM UNI F 1 <1 PO EX

PAC-09BO-033-02 B 4 4 UNIM UNI F 3 2–3 PO EX, RT

PAC-09BO-033-04 B 4 4 ODO CAL DE+DS L 1 FU 2.9 1.7 PO EX, SH, CRK PAC-09BO-034-01 B 4 4 UNI UNI F 2 <1 PO EX, RT

PAC-09BO-034-02 B 4 4 UNIM UNI F 3 1–2 PO EX, RT

PAC-09BO-034-03 B 4 4 UNIM UNI F 2 1–2 SD EX

PAC-09BO-034-04 B 4 4 UNIM UNI F 1 1–2 PO EX, CRK, BUR

PAC-09BO-034-05 B 4 4 UNIM LBF F 1 2.6 1.1 PO EX, RT

PAC-09BO-034-06 B 4 4 STM-MTM LBF F 1 1.2 0.7 DA CRK, BUR

PAC-09BO-034-07 B 4 4 MTM-LTM MO-PM F 1 1.5 0.6 PO EX, RT

PAC-09BO-035-01 B 4 4 UNI UNI F 4 <1 PO EX

PAC-09BO-035-02 B 4 4 UNIM UNI F 4 1–2 PO EX, RT

PAC-09BO-035-03 B 4 4 UNIM UNI F 1 3–4 PO EX, RT

PAC-09BO-036-01 B 4 4 ODO MO UP L 1 1.1 0.6 SD EX, BUR

PAC-09BO-037-01 B 4 4 UNIM UNI F 11 <1 PO EX

PAC-09BO-037-02 B 4 4 UNIM UNI F 2 1–2 PO EX

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-09BO-037-04 B 4 4 UNIM UNI F 2 2–3 PO EX, RT

PAC-09BO-037-05 B 4 4 ODO RAD PE R 1 FU 1.4 1.0 SD EX PAC-09BO-037-06 B 4 4 MTM VER F 1 1.5 0.7 PO EX, RT

PAC-09BO-038-01 B 4 5 UNIM UNI F 1 1–2 PO EX

PAC-09BO-039-01 B 4 5 UNIM UNI F 3 1–2 PO EX

PAC-09BO-039-02 B 4 5 MTM-LTM LBF F 1 3.0 1.3 PO EX, RT

PAC-09BO-039-03 B 4 5 ODO MTT PE L 1 FU 2.1 1.5 DA EX PAC-09BO-040-01 B 4 5 UNI UNI F 3 <1 PO EX

PAC-09BO-040-02 B 4 5 UNIM UNI F 9 1–2 PO EX

PAC-09BO-040-03 B 4 5 UNIM UNI F 3 2–3 PO EX

PAC-09BO-040-04 B 4 5 UNIM LBF F 1 2.6 1.3 DA EX

PAC-09BO-040-06 B 4 5 ODO GC F R 1 FU 1.4 0.9 PO EX PAC-09BO-040-07 B 4 5 ODO RAD DE R 1 4.0 1.1 PO EX, RT

PAC-09BO-040-08 B 4 5 MTM-LTM LBF F 1 5.1 1.2 PO EX, RT

PAC-09BO-042-01 B 4 5 UNIM LBF F 1 2.7 1.1 PO EX

PAC-09BO-043-01 B 5 4 UNIM UNI F 1 1–2 PO EX

PAC-09BO-044-01 B 5 4 UNIM UNI F 1 1–2 PO EX

PAC-09BO-049-01 B 5 4 UNIM UNI F 1 1–2 SD EX

PAC-09BO-050-01 B 5 4 UNIM UNI F 1 1–2 PO EX

PAC-09BO-050-02 B 5 4 UNIM UNI F 1 1–2 PO EX

PAC-09BO-051-01 B 6 4 UNIM UNI F 1 1–2 PO EX, RT

PAC-09BO-052-01 B 6 4 LTM LBF F 1 2.9 1.6 PO EX, RT

PAC-09BO-053-01 B 7 4 MTM-LTM TTH F 1 1.3 0.7 PO RT

PAC-09BO-054-01 B 8 4 UNIM UNI F 1 <1 PO EX

PAC-09BO-054-02 B 8 4 UNIM UNI F 3 1–2 PO EX

PAC-09BO-054-03 B 8 4 UNIM UNI F 1 2–3 PO EX

PAC-09BO-054-04 B 8 4 UNIM UNI F 1 1–2 PO EX

229

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-09BO-055-01 B 10 4 UNIM UNI F 4 <1 PO EX

PAC-09BO-055-02 B 10 4 UNIM UNI F 4 1–2 PO EX

PAC-09BO-056-01 B 2 4 UNI UNI F 5 <1 PO EX

PAC-09BO-056-02 B 2 4 UNIM UNI F 13 1–2 PO EX

PAC-09BO-056-03 B 2 4 UNIM UNI F 5 1–2 DA EX

PAC-09BO-056-04 B 2 4 UNIM UNI F 2 2–3 PO EX, RT

PAC-09BO-056-05 B 2 4 UNIM LBF F 1 1.3 1.2 SD EX, RT, BUR

PAC-09BO-056-08 B 2 4 MTM-LTM RIB F 1 3.8 0.8 PO EX, SH

PAC-09BO-056-09 B 2 4 ODO MO2 LO R 1 YEA 1.5 1.5 SD EX, RT PAC-09BO-057-01 B 2 4 UNIM UNI F 2 1–2 PO EX, RT

PAC-09BO-058-01 B 4 4 UNI UNI F 11 <1 PO EX

PAC-09BO-058-02 B 4 4 UNIM UNI F 9 1–2 PO EX

PAC-09BO-058-03 B 4 4 UNIM UNI F 3 1–2 PO EX

PAC-09BO-058-04 B 4 4 LTM LBF F 1 5.9 1.9 PO EX

PAC-09BO-058-05 B 4 4 ARM SHL F 1 0.8 0.6 DA EX

PAC-09BO-058-06 B 4 4 TAY PH1 PE 1 U 0.9 0.8 PO EX

PAC-09BO-058-07 B 4 4 ODO MAL F L 1 FU 1.6 1.3 PO EX, RT PAC-09BO-058-08 B 4 4 ODO DP4 LO R 1 YEA 1.4 0.9 PO EX PAC-09BO-058-09 B 4 4 CER MO-PM UP 1 1.1 0.6 DA EX, RT

PAC-09BO-059-01 B 4 5 UNIM UNI F 1 <1 PO EX

PAC-09BO-059-02 B 4 5 UNIM UNI F 3 1–2 PO EX

PAC-09BO-059-03 B 4 5 MTM-LTM LBF F 1 5.7 1.3 DA EX, RT, CRK, GN

PAC-09BO-059-04 B 4 5 LTM LBF F 1 7.8 1.7 PO EX, RT

PAC-09BO-059-05 B 4 5 ODO MTC PE+PS L 1 FU 6.7 2.4 PO EX, RT PAC-09BO-059-06 B 4 5 ODO AST W R 1 FU 3.6 2.3 DA EX, RT, GN PAC-09BO-059-07 B 4 5 ODO MTC DS L 1 5.6 1.3 PO EX, RT, CRK, STG

PAC-09BO-059-08 B 4 5 GAL FIB PE R 1 U 1.5 0.8 DA EX, RT, STG

230

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-09BO-060-01 B 10 4 DOG HUM PS L 1 2.5 1.2 PO EX

PAC-09BO-061-01 B 10 4 UNIM UNI F 1 1–2 PO EX

PAC-09BO-062-01 B 10 4 UNIM UNI F 1 1–2 PO EX

PAC-09BO-062-02 B 10 4 ODO TIB DS R 1 4.3 2.0 PO EX, RT

PAC-09BO-063-01 B 10 4 UNI UNI F 1 <1 PO EX

PAC-09BO-063-02 B 10 4 UNIM UNI F 2 1–2 PO EX

PAC-09BO-063-03 B 10 4 UNI UNI F 1 1.3 0.4 PO BUR

PAC-09BO-063-04 B 10 4 UNIM LBF F 1 1.6 1.1 PO EX, RT

PAC-09BO-063-05 B 10 4 PAF PMX PRO L 1 1.8 0.9 SD EX

PAC-09BO-063-06 B 10 4 ODO NVB F L 1 FU 1.8 1.4 PO EX PAC-09BO-064-01 B 10 4 UNIM UNI F 3 1–2 PO EX

PAC-09BO-064-02 B 10 4 UNIM LBF F 1 2.7 0.9 PO EX, RT

PAC-10BO-006-01 A 1 5 GOP FEM PE+PS+DE R 1 FU 2.3 1.3 SD

PAC-10BO-007-01 A 1 5 UNIM UNI F 1 3–4 PO EX, RT

PAC-10BO-014-01 B 3 5 UNIM UNI F 6 1–2 DA EX

PAC-10BO-014-03 B 3 5 ODO MAL F L 1 FU 1.5 1.1 SD EX PAC-10BO-014-04 B 3 5 IGU CDV F AX 1 1.1 0.7 SD EX

PAC-10BO-014-05 B 3 5 MAZ PH1 DE+S 1 FU 2.0 0.7 DA SH, CRK

PAC-10BO-014-06 B 3 5 MUS MAN+LM2 F R 1 1.8 0.8 SD EX

PAC-10BO-014-07 B 3 5 ODO MO3 UP L 1 1.5 1.2 SD EX, RT

PAC-10BO-014-08 B 3 5 UNIM UNI F 1 <1 DA EX

PAC-10BO-020-01 B 5 5 UNI UNI F 1 <1 PO EX

PAC-10BO-020-02 B 5 5 UNIM UNI F 3 1–2 DA EX, STG

PAC-10BO-020-03 B 5 5 UNIM UNI F 2 2–3 PO EX

PAC-10BO-020-04 B 5 5 UNIM UNI F 1 1.2 0.8 SD RT, BUR

PAC-10BO-020-05 B 5 5 LTM LBF F 1 5.4 1.2 PO EX, RT

PAC-10BO-020-08 B 5 5 TAY FEM PE L 1 FU 1.4 0.7 DA EX

231

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-10BO-020-09 B 5 5 ODO RIB F L 1 1.8 1.4 DA EX

PAC-10BO-020-10 B 5 5 ODO MO UP 1 1.1 0.9 PO EX, RT

PAC-10BO-020-11 B 5 5 ODO MO-PM UP R 1 1.0 0.7 SD EX

PAC-10BO-029-01 B 6 5 ODO TIB DS+DE L 1 FU 4.1 2.2 SD EX, RT, STG PAC-10BO-029-02 B 6 5 RAB INN ACE L 1 FU 1.9 0.9 PO EX, RT, STG PAC-10BO-030-01 B 6 5 UNI UNI F 2 <1 PO EX

PAC-10BO-030-02 B 6 5 UNIM UNI F 1 1–2 PO EX, RT

PAC-10BO-030-03 B 6 5 MAZ NVB W R 1 FU 2.0 1.5 DA EX, RT PAC-10BO-031-01 B 6 5 UNI UNI F 4 <1 PO EX

PAC-10BO-031-02 B 6 5 UNIM UNI F 4 1–2 PO EX

PAC-10BO-031-03 B 6 5 UNIM UNI F 2 2–3 PO EX, RT

PAC-10BO-031-04 B 6 5 MTM-LTM LBF F 1 4.4 0.8 PO EX, RT

PAC-10BO-031-05 B 6 5 LTM LBF F 1 3.6 1.6 PO EX

PAC-10BO-031-06 B 6 5 ODO MO3 LO R 1 ADU 2.3 0.9 DA EX PAC-10BO-031-07 B 6 5 ODO MO1-2 LO R 1 YEA 2.3 1.0 DA EX PAC-10BO-037-01 B 8 5 ODO PH2 PE+PS 1 2.0 1.5 PO EX

PAC-10BO-039-01 B 9 5 VIP VER F AX 1 1.0 0.9 PO EX

PAC-96BO-065-01 B 2b 3 UNIM UNI F 2 2–3 PO EX, RT

PAC-96BO-065-02 B 2b 3 UNIM UNI F 1 3.2 0.9 PO EX, RT, BUR

PAC-96BO-065-03 B 2b 3 ODO PH1 DE 1 FU 1.7 1.0 PO EX

PAC-96BO-069-01 B 3c 4 UNI UNI F 1 <1 PO EX

PAC-96BO-069-02 B 3c 4 UNIM UNI F 1 1–2 PO EX

PAC-96BO-069-03 B 3c 4 UNIM UNI F 2 1–2 PO EX

PAC-96BO-069-05 B 3c 4 MTM-LTM LBF F 1 3.2 1.6 PO EX, RT

PAC-96BO-069-06 B 3c 4 LTM LBF F 1 5.9 2.1 PO EX, RT

PAC-96BO-073-01 B 1 3 UNIM UNI F 1 2–3 PO EX, RT

PAC-96BO-074-01 B 8 3 UNI UNI F 3 <1 PO EX

232

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-96BO-074-02 B 8 3 UNIM UNI F 9 1–2 PO EX, RT

PAC-96BO-074-03 B 8 3 UNIM UNI F 2 2–3 PO EX, RT

PAC-96BO-074-07 B 8 3 MTM-LTM LBF F 1 3.9 1.2 DA EX, RT

PAC-96BO-074-10 B 8 3 ODO FEM PS L 1 3.5 1.9 PO EX

PAC-96BO-077-01 B 3 3 UNIM UNI F 1 3–4 SD EX, BUR

PAC-96BO-078-01 B 3b 3 LTM UNI F 1 3.5 1.6 PO EX, RT

PAC-96BO-079-01 B 11 3 UNI UNI F 1 <1 DA EX

PAC-96BO-079-02 B 11 3 STM LBF F 1 1.5 0.3 PO EX, RT

PAC-96BO-080-01 B 6 3 UNIM UNI F 1 <1 PO EX

PAC-96BO-080-02 B 6 3 MAZ MTT DS 1 1.8 0.8 PO EX

PAC-96BO-080-03 B 6 3 ODO FEM DE R 1 FU 2.8 1.8 PO EX, CRK PAC-96BO-080-04 B 6 3 ARM PH1 W L 1 FU 0.6 0.5 DA EX PAC-96BO-080-05 B 6 3 COL VER W AX 1 1.0 0.9 PO EX

PAC-96BO-080-06 B 6 3 COL VER W AX 1 1.0 0.9 PO EX

PAC-96BO-080-07 B 6 3 COL VER W AX 1 1.0 1.0 PO EX

PAC-96BO-081-01 B 3 5 UNI UNI F 3 <1 PO EX, RT

PAC-96BO-081-02 B 3 5 UNIM UNI F 16 1–2 PO EX, RT

PAC-96BO-081-03 B 3 5 UNIM UNI F 5 2–3 PO EX

PAC-96BO-081-04 B 3 5 MTM-LTM LBF F 1 2.9 1.2 PO EX, RT

PAC-96BO-081-05 B 3 5 MTM-LTM LBF F 1 2.9 1.1 PO EX, RT

PAC-96BO-081-07 B 3 5 ODO MTC PE+PS+MS L 1 FU 11.1 2.2 SD EX, RT, GN PAC-96BO-081-08 B 3 5 ODO LMV F AX 1 FU 1.0 0.9 PO EX PAC-96BO-081-09 B 3 5 ODO PH2 PE 1 FU 1.2 1.0 SD

PAC-96BO-081-10 B 3 5 LTM LBF F 1 5.4 1.8 DA EX, RT

PAC-96BO-082-01 B 3d 2 ARM TIB MS+DS L 1 2.5 0.8 PO EX, RT, STG

PAC-96BO-083-01 B 6 5 UNI UNI F 3 <1 PO EX

PAC-96BO-083-02 B 6 5 UNIM UNI F 4 1–2 PO EX

233

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-96BO-083-03 B 6 5 UNIM UNI F 1 3–4 SD EX, RT

PAC-96BO-083-05 B 6 5 ODO MTT PE L 1 FU 2.0 1.7 DA EX PAC-96BO-083-06 B 6 5 ODO DP4 LO L 1 FAW 1.5 0.7 SD EX PAC-96BO-083-07 B 6 5 ODO MO1 LO L 1 YEA 1.9 0.8 DA EX PAC-96BO-085-01 B 4 3 UNIM UNI F 1 2–3 PO EX, RT

PAC-96BO-086-01 B 5 3 UNIM UNI F 1 2–3 PO EX, RT

PAC-96BO-087-01 B 6 2 UNIM UNI F 1 3–4 PO EX, RT

PAC-96BO-088-01 B 3b 2 UNIM UNI F 1 1.6 0.9 SD RT, BUR

PAC-96BO-089-01 B 9 3 ODO FEM DS L 1 5.3 2.3 DA EX

PAC-96BO-090-01 B 3b 4 UNI UNI F 1 <1 PO EX

PAC-96BO-090-02 B 3b 4 UNI UNI F 2 1–2 PO EX

PAC-96BO-090-03 B 3b 4 UNIM UNI F 2 1–2 DA EX

PAC-96BO-090-04 B 3b 4 UNIM UNI F 1 3–4 PO EX, RT

PAC-96BO-090-06 B 3b 4 LTM LBF F 1 3.9 1.5 PO EX, RT

PAC-96BO-091-01 B 2d 4 UNIM UNI F 2 1–2 DA EX

PAC-96BO-091-03 B 2d 4 ARM MTC DE+DS 1 FU 0.7 0.5 DA EX

PAC-96BO-092-01 B 6 3 MTM-LTM LBF F 1 4.2 1.2 PO EX, RT

PAC-96BO-093-01 B 5 4 KIN SHL F 1 1.4 1.0 PO EX

PAC-96BO-094-01 B 6 5 UNI UNI F 9 <1 DA EX

PAC-96BO-094-02 B 6 5 UNIM UNI F 3 1–2 DA EX, RT

PAC-96BO-094-03 B 6 5 UNIM UNI F 1 2–3 PO EX

PAC-96BO-094-04 B 6 5 UNIM UNI F 1 2–3 PO EX, RT

PAC-96BO-094-06 B 6 5 ARM SHL F 1 0.5 0.5 DA EX

PAC-96BO-094-07 B 6 5 CER MO-PM F 1 1.0 0.5 SD RT

PAC-96BO-095-01 B 13 5 UNIM UNI F 1 1–2 PO EX

PAC-96BO-095-02 B 13 5 UNIM UNI F 1 2–3 PO EX

PAC-96BO-096-01 B 2d 5 UNI UNI F 1 <1 PO EX

234

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-96BO-096-02 B 2d 5 UNIM UNI F 4 1–2 DA EX

PAC-96BO-097-01 B 6 5 UNIM UNI F 1 2–3 SD EX

PAC-96BO-098-01 B 7 5+6 UNIM UNI F 1 2–3 DA EX

PAC-96BO-098-02 B 7 5+6 ODO MTT DS R 1 3.7 1.7 PO EX, CRK

PAC-96BO-098-03 B 7 5+6 ODO RIB POST R 1 FU 2.4 1.3 SD EX PAC-96BO-101-01 B 5 5 UNI UNI F 3 <1 DA EX

PAC-96BO-101-02 B 5 5 UNIM UNI F 5 1–2 DA EX

PAC-96BO-101-03 B 5 5 UNIM LBF F 1 2–3 SD EX

PAC-96BO-101-04 B 5 5 ARM SHL F 1 0.7 0.5 DA EX

PAC-96BO-101-05 B 5 5 ODO GC F R 1 U 1.7 0.9 SD EX PAC-96BO-101-07 B 5 5 LTM LBF F 1 6.5 1.2 PO EX, RT, SH

PAC-96BO-102-01 B 11 3 UNI UNI F 1 1–2 PO EX, RT

PAC-96BO-104-01 B 10 4 UNIM UNI F 1 2–3 PO EX

PAC-96BO-104-02 B 10 4 UNIM UNI F 1 3–4 PO EX

PAC-96BO-104-03 B 10 4 MTM-LTM LBF F 1 2.3 1.0 PO EX

PAC-96BO-104-04 B 10 4 ODO TRQ W R 1 FU 2.1 1.4 PO EX PAC-96BO-104-05 B 10 4 ODO MAL W L 1 FU 1.6 1.5 SD EX PAC-96BO-106-01 B 2c 5 UNIM UNI F 1 1–2 PO EX

PAC-96BO-106-02 B 2c 5 UNIM UNI F 2 1–2 PO EX

PAC-96BO-106-03 B 2c 5 ARM SHL F 1 0.7 0.5 SD EX

PAC-96BO-106-04 B 2c 5 ODO MTT PE R 1 FU 2.0 0.8 DA EX PAC-96BO-122-01 B 1 5 UNI UNI F 1 <1 PO EX

PAC-96BO-122-02 B 1 5 UNIM UNI F 2 1–2 PO EX

PAC-96BO-123-01 B 3d 5 UNIM UNI F 1 2–3 PO EX, CRK

PAC-96BO-123-02 B 3d 5 LTM LBF F 1 2.9 1.5 DA EX, RT, PN

PAC-96BO-123-03 B 3d 5 LTM LBF F 1 3.6 1.6 PO EX

PAC-96BO-123-04 B 3d 5 MTM-LTM LBF F 1 2.9 0.7 DA SH, CRK, STG

235

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-96BO-124-01 B 3 5 ARM SHL F 1 0.6 0.5 IT BUR

PAC-96BO-125-01 B 3 5 UNI UNI F 3 <1 DA EX

PAC-96BO-125-02 B 3 5 UNIM UNI F 3 1–2 PO EX

PAC-96BO-125-03 B 3 5 UNIM LBF F 1 2.9 1.0 PO EX

PAC-96BO-126-01 B 2b 5 UNIM UNI F 1 1–2 PO EX

PAC-96BO-127-01 B 7 5+6 UNIM UNI F 3 1–2 PO EX

PAC-96BO-127-02 B 7 5+6 UNIM UNI F 1 4–5 DA EX

PAC-96BO-127-03 B 7 5+6 LTM LBF F 1 6.8 1.7 SD EX, RT

PAC-96BO-128-01 B 4 5 UNIM LBF F 1 2–3 PO EX

PAC-96BO-128-02 B 4 5 ODO FEM DE L 1 FU 2.2 2.0 PO EX PAC-96BO-129-01 B 8 5 UNI UNI F 2 <1 DA EX

PAC-96BO-129-02 B 8 5 UNIM UNI F 6 1–2 PO EX

PAC-96BO-129-03 B 8 5 UNIM UNI F 1 3–4 PO EX, CRK

PAC-96BO-129-04 B 8 5 OPOV MAN F L 1 1.3 0.9 PO EX

PAC-96BO-129-05 B 8 5 ARM RAD PE+PS+MS R 1 2.0 0.4 PO EX, RT

PAC-96BO-129-06 B 8 5 ODO MO UP R 1 1.5 1.0 PO EX

PAC-96BO-129-07 B 8 5 STM LBF F 1 1.2 0.5 DA SH

PAC-96BO-130-01 B 12 5 UNIM UNI F 1 2–3 DA EX

PAC-96BO-130-02 B 12 5 LTM LBF F 1 3.6 1.7 PO EX, SH

PAC-96BO-130-03 B 12 5 LTM UNI F 1 3.0 1.7 PO EX

PAC-96BO-131-01 B 10 5 STM LBF F 1 2.2 0.5 DA EX, GN

PAC-96BO-131-02 B 10 5 CAT PH1 DE+DS 1 FU 0.6 0.6 SD

PAC-96BO-133-01 B 7 5 UNIM UNI F 1 3–4 PO EX, RT

PAC-96BO-134/141-01 B 1 5 ODO HUM DS R 1 5.2 1.7 PO EX, RT, SH

PAC-96BO-137-01 B 11 3 UNIM UNI F 1 1–2 PO EX

PAC-96BO-138-01 B 1 3 UNIM UNI F 2 <1 PO EX

PAC-96BO-138-02 B 1 3 UNIM UNI F 1 1–2 PO EX, CRK

236

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-96BO-139-01 B 1 4 UNI UNI F 1 <1 PO EX

PAC-96BO-140-01 B 1 5 UNIM UNI F 3 1–2 DA EX

PAC-96BO-140-02 B 1 5 UNIM UNI F 1 1–2 PO EX

PAC-96BO-140-03 B 1 5 LTM LBF F 1 4.8 1.2 PO EX, SH, CRK

PAC-96BO-141-01 B 1 5 UNI UNI F 2 <1 SD EX

PAC-96BO-141-02 B 1 5 UNI UNI F 1 <1 PO EX

PAC-96BO-141-03 B 1 5 UNIM UNI F 1 <1 PO EX

PAC-96BO-141-04 B 1 5 UNIM UNI F 1 1–2 DA EX, RT

PAC-96BO-141-05 B 1 5 UNIM UNI F 1 1–2 PO EX

PAC-96BO-144-01 B 2b 4 UNIM UNI F 1 <1 PO EX

PAC-96BO-145-01 B 3b 4 UNIM UNI F 1 1–2 PO EX

PAC-96BO-146-01 B 3 5 UNI UNI F 1 <1 PO EX

PAC-96BO-150-01 B 5 5 UNIM UNI F 1 1–2 PO EX, RT

PAC-96BO-150-02 B 5 5 STM-MTM LBF F 1 2.3 0.7 SD RT

PAC-96BO-152-01 B 6 5 UNIM UNI F 1 1–2 PO EX

PAC-96BO-152-02 B 6 5 ODO MTP S 1 1.3 0.7 SD RT, STG

PAC-96BO-152-03 B 6 5 TES SHL F 1 0.8 0.6 DA EX

PAC-96BO-153-01 B 7 3+5 UNIM UNI F 2 1–2 PO EX

PAC-96BO-154-01 B 7 3 UNIM UNI F 1 2–3 PO EX

PAC-96BO-155-01 B 8 3 UNI UNI F 1 <1 PO EX

PAC-96BO-155-02 B 8 3 UNIM UNI F 8 1–2 PO EX

PAC-96BO-155-04 B 8 3 UNIM UNI F 1 2–3 PO EX

PAC-96BO-155-05 B 8 3 TAY PH2 W 1 FU 1.6 1.0 PO EX, RT, GN

PAC-96BO-155-06 B 8 3 MAZ HUM DE L 1 FU 1.5 1.3 PO EX PAC-96BO-155-07 B 8 3 MTM-LTM LBF F 1 4.9 0.6 PO EX, RT

PAC-96BO-156-01 B 8 5/6B UNI UNI F 4 <1 DA EX

PAC-96BO-156-02 B 8 5/6B UNI UNI F 3 1–2 DA EX

237

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-96BO-156-03 B 8 5/6B UNI UNI F 1 2–3 DA EX, RT

PAC-96BO-156-04 B 8 5/6B TAY NAV W R 1 FU 1.7 1.1 PO EX, RT PAC-96BO-159-01 B 9 5 UNI UNI F 6 <1 PO EX

PAC-96BO-159-02 B 9 5 UNIM UNI F 2 1–2 PO EX

PAC-96BO-159-03 B 9 5 ODO PH2 W 1 FU 3.0 1.3 PO EX, GN

PAC-96BO-159-05 B 9 5 TAY PH2 W 1 FU 1.7 1.0 PO EX

PAC-97BO-002-01 B 97-U5 3 UNI UNI F 10 <1 PO

PAC-97BO-002-02 B 97-U5 3 UNIM UNI F 9 <1 PO EX

PAC-97BO-002-03 B 97-U5 3 UNI UNI F 1 1–2 PO

PAC-97BO-002-04 B 97-U5 3 UNIM UNI F 1 2–3 PO

PAC-97BO-002-05 B 97-U5 3 UNIM UNI F 1 1–2 PO EX

PAC-97BO-002-06 B 97-U5 3 UNIM UNI F 1 2–3 PO EX

PAC-97BO-002-07 B 97-U5 3 UNIM UNI F 1 2–3 SD EX

PAC-97BO-002-08 B 97-U5 3 ODO FEM DE L 1 FU 5.5 5.3 DA EX, RT, CT, GN PAC-97BO-002-09 B 97-U5 3 ODO FEM PE L 1 FU 4.5 3.1 DA EX, RT, GN PAC-97BO-002-10 B 97-U5 3 ODO FEM PS+MS+DS L 1 21.2 4.8 DA EX, RT, CRK

PAC-97BO-002-11 B 97-U5 3 ODO CAL DE+DS+MS R 1 FU 6.3 2.3 DA EX PAC-97BO-002-12 B 97-U5 3 ODO HUM DS L 1 5.5 1.8 SD EX

PAC-97BO-002-13 B 97-U5 3 ODO AST F R 1 FU 1.6 1.5 SD EX PAC-97BO-003-01 B 97-U2 3 UNIM UNI F 1 <1 PO EX

PAC-97BO-003-02 B 97-U2 3 UNI UNI F 1 <1 PO EX

PAC-97BO-003-03 B 97-U2 3 UNIM UNI F 1 1–2 PO EX

PAC-97BO-004-01 B 97-SU1 3 UNIM UNI F 2 1–2 PO EX

PAC-97BO-004-02 B 97-SU1 3 UNIM UNI F 1 1–2 DA EX

PAC-97BO-004-04 B 97-SU1 3 UNIM UNI F 1 2–3 PO EX, RT

PAC-97BO-005-01 B 97-U4 3 UNI UNI F 1 <1 PO EX

PAC-97BO-005-02 B 97-U4 3 UNI UNI F 2 1–2 PO EX

238

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-97BO-006-01 C 97C5 4 UNIM UNI F 4 <1 PO EX

PAC-97BO-006-02 C 97C5 4 UNIM UNI F 4 <1 PO EX

PAC-97BO-006-03 C 97C5 4 UNIM UNI F 1 1–2 SD EX

PAC-97BO-006-04 C 97C5 4 UNIM UNI F 2 1–2 DA EX, CRK

PAC-97BO-006-05 C 97C5 4 UNIM UNI F 1 1–2 PO EX

PAC-97BO-006-06 C 97C5 4 LTM UNI F 1 3.3 1.1 PO EX, CRK

PAC-97BO-009-01 C 97C2 4 UNIM UNI F 1 3–4 DA EX

PAC-97BO-010-01 B 97-RU1 5 UNIM UNI F 1 1–2 DA EX

PAC-97BO-010-02 B 97-RU1 5 UNIM UNI F 1 1–2 SD EX

PAC-97BO-010-03 B 97-RU1 5 UNIM UNI F 1 2–3 PO EX

PAC-97BO-012-01 B 97-RU1 3 UNIM UNI F 1 1–2 PO EX

PAC-97BO-012-02 B 97-RU1 3 UNIM UNI F 1 2–3 PO EX

PAC-97BO-013-01 B 97-SU4 3 UNIM UNI F 1 1–2 PO EX

PAC-97BO-019-01 B 97-U3 3 UNIM UNI F 1 <1 PO EX

PAC-97BO-019-02 B 97-U3 3 UNIM UNI F 3 1–2 PO EX

PAC-97BO-019-03 B 97-U3 3 UNIM UNI F 1 1–2 PO EX

PAC-97BO-019-04 B 97-U3 3 UNIM UNI F 2 1–2 SD EX

PAC-97BO-019-05 B 97-U3 3 UNIM UNI F 1 2–3 DA EX

PAC-97BO-019-06 B 97-U3 3 ARM SHL F 1 0.7 0.6 PO EX

PAC-97BO-020-02 C 97C5 5 UNIM UNI F 1 1.9 1.4 DA EX, RT

PAC-97BO-020-03 C 97C5 5 UNIM UNI F 1 1–2 PO EX

PAC-97BO-020-04 C 97C5 5 UNIM UNI F 1 2–3 PO EX

PAC-97BO-023-01 B 97-U4 5 UNI UNI F 5 <1 PO EX

PAC-97BO-023-02 B 97-U4 5 UNIM UNI F 9 1–2 PO EX

PAC-97BO-023-03 B 97-U4 5 UNIM UNI F 1 1–2 SD EX, RT, STG

PAC-97BO-023-04 B 97-U4 5 UNIM UNI F 1 2–3 PO EX, RT

PAC-97BO-023-06 B 97-U4 5 ODO RIB F R 1 FU 2.0 0.9 DA EX

239

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-97BO-023-07 B 97-U4 5 TES SHL F 1 1.0 0.8 PO EX, RT

PAC-97BO-024-01 B 97-TU4 5 UNIM UNI F 1 1–2 IT

PAC-97BO-024-02 B 97-TU4 5 UNIM UNI F 1 1–2 PO EX

PAC-97BO-028-01 B 97-TU2 3 UNIM UNI F 1 <1 DA EX

PAC-97BO-028-02 B 97-TU2 3 UNIM UNI F 1 1–2 DA EX

PAC-97BO-028-03 B 97-TU2 3 UNIM UNI F 1 3–4 SD EX

PAC-97BO-032-01 B 97-SU5 4 UNIM UNI F 2 1–2 PO EX

PAC-97BO-032-02 B 97-SU5 4 UNIM UNI F 1 2–3 PO EX

PAC-97BO-032-03 B 97-SU5 4 UNIM UNI F 1 3–4 PO EX

PAC-97BO-033-01 B 97-SU5 3 UNIM UNI F 3 <1 PO EX

PAC-97BO-033-02 B 97-SU5 3 UNIM UNI F 3 1–2 PO EX

PAC-97BO-036-01 B 97-TU1 3 UNIM UNI F 1 <1 PO EX

PAC-97BO-036-02 B 97-TU1 3 UNIM UNI F 1 1–2 PO EX

PAC-97BO-036-03 B 97-TU1 3 UNIM UNI F 1 1–2 SD EX

PAC-97BO-036-04 B 97-TU1 3 UNIM UNI F 1 2–3 DA EX

PAC-97BO-039-01 B 97-U4 5 UNI UNI F 5 <1 PO EX

PAC-97BO-039-02 B 97-U4 5 UNIM UNI F 3 1–2 SD EX

PAC-97BO-039-03 B 97-U4 5 UNIM UNI F 2 1–2 DA EX, RT, CRK

PAC-97BO-039-04 B 97-U4 5 UNIM UNI F 5 2–3 PO EX, RT

PAC-97BO-039-05 B 97-U4 5 UNIM UNI F 1 3–4 PO EX

PAC-97BO-039-06 B 97-U4 5 UNIM UNI F 1 2.0 1.4 PO RT, BUR

PAC-97BO-039-12 B 97-U4 5 MTM MAX F R 1 2.2 1.7 DA EX

PAC-97BO-039-13 B 97-U4 5 ARM TIB DS+MS L 1 3.3 0.9 PO EX, STG

PAC-97BO-040-01 B 97-U4 4 UNIM UNI F 1 2–3 PO EX

PAC-97BO-040-02 B 97-U4 4 OPOM FEM PS+MS+DS R 1 4.2 0.6 PO EX, RT, STG

PAC-97BO-041-01 B 97-U4 4 UNIM UNI F 1 1–2 PO EX, CRK

PAC-97BO-041-02 B 97-U4 4 UNIM UNI F 1 3–4 PO EX, RT

240

Max Max Surface Catalogue number Plaza Unit Level ID Bone End Side Count Fusion Taphonomy length width state PAC-97BO-041-03 B 97-U4 4 LTM LBF F 1 4.0 1.3 SD RT

PAC-97BO-041-04 B 97-U4 4 LTM UNI F 1 2.8 1.0 PO EX

PAC-97BO-043-01 B 97-U5 3 ODO HUM DS R 1 3.1 2.1 PO EX, CRK

PAC-97BO-043-02 B 97-U5 3 ODO HUM DE+DS R 1 FU 4.9 2.4 PO EX, RT, GN PAC-97BO-043-03 B 97-U5 3 MTM-LTM UNI F 1 2.5 0.7 PO EX, RT, CRK, STG

PAC-97BO-043-04 B 97-U5 3 MTM-LTM UNI F 1 2.2 0.4 PO RT, CRK, STG

PAC-97BO-045-01 B 97-SU9 4 RAB NAV W L 1 FU 0.8 0.6 PO EX PAC-97BO-050-01 B 97-U1 5 LTM LBF F 1 3.6 1.5 PO EX, RT

PAC-97BO-054-01 B 97-TU2 5 UNIM UNI F 1 <1 PO EX

PAC-97BO-055-01 B 97-SU4 4 UNIM LBF F 1 1.0 0.5 IT BUR

PAC-97BO-057-01 B 97-SU2 3 UNIM UNI F 1 <1 DA EX

PAC-97BO-058-01 B 97-SU2 4 UNIM UNI F 1 1–2 PO EX

PAC-97BO-060-01 C 97C1 3 UNIM UNI F 1 1–2 PO EX, RT

PAC-97BO-062-01 B 97-SU3 4 UNIM UNI F 1 1–2 DA EX

PAC-97BO-063-01 B 97-SU2 3 UNIM UNI F 7 <1 SD EX

PAC-97BO-063-02 B 97-SU2 3 UNIM UNI F 5 1–2 PO EX

PAC-97BO-063-03 B 97-SU2 3 UNIM UNI F 1 1–2 SD EX, RT

PAC-97BO-063-04 B 97-SU2 3 UNIM UNI F 1 2–3 DA EX, CRK

PAC-97BO-133-01 B 97-SU2 3 UNI UNI F 1 <1 DA EX

241

Table A.3 Pacbitun dataset for the transitional period late Middle Preclassic-early Late Preclassic

Catalogue Max Max Surface Plaza Unit Level ID Bone End Side Count Fusion Taphonomy number length width state PAC-10BO-012-01 A 2 4 UNI UNI F 1 1–2 PO EX

PAC-10BO-012-02 A 2 4 MAZ MTT PE+PS+MS L 1 FU 6.6 1.2 PO EX, RT, CRK PAC-10BO-012-03 A 2 4 ODO PH2 W 1 FU 3.3 1.9 DA EX, RT, GN

PAC-10BO-013-01 A 2 4 UNI UNI F 1 <1 PO EX

PAC-10BO-013-02 A 2 4 ODO PH2 W 1 FU 2.6 1.4 PO EX

242