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An Investigation of Late Upper Paleolithic and Epipaleolithic Hunter-

Gatherer Subsistence and Settlement Patterns in Central Portugal


Jonathan Adams Haws

A dissertation submitted in partial fulfillment of the requirements for

the degree of

Doctor of Philosophy


at the


An Investigation of Late Upper Paleolithic and Epipaleolithic Hunter-Gatherer
Subsistence and Settlement Patterns in Central Portugal
Jonathan Adams Haws
Under the supervision of Professor T. Douglas Price
At the University of Wisconsin-Madison

This study investigates the nature of Upper Paleolithic hunter-gatherer subsistence

and settlement patterns in central Portugal. The primary research goal is to test the Broad

Spectrum Revolution model which is the predominate explanatory framework used by

archaeologists in the region. Previous models characterized evidence of diachronic changes

in subsistence as an indication of increased resource intensification, specialization and

diversification during the Late Upper Paleolithic. In Iberia, archaeologists characterize

intensification as the extraction of greater amounts of energy from the same resource.

Specialization is seen through the occurrence of sites whose function centers around specific

tasks and the focus on limited numbers of resource types.

The goal of this dissertation is to show that the main components of the Broad Spectrum

Revolution model, resource intensification and diversification, did not suddenly appear

at the beginning of the Holocene, but that they have a much greater time depth. It is

argued here that dietary diversity is part of our evolutionary heritage as omnivorous

primates and shifts between generalized and specialized diets reflect local climatic and

environmental conditions, not a directional trend in human adaptation.

To test this model, archaeofaunal assemblages from two Late Upper Paleolithic caves

in central Portugal were analyzed. In addition, plant exploitation was addressed by using

the regional archaeological record, paleoenvironmental reconstruction and expectations

from evolutionary ecology. The assemblages from Lapa do Picareiro and Lapa do Suão

represent the best samples that date to the Late Pleistocene and Early Holocene. Each site

has multicomponent occupations allowing the study of diachronic trends in resource use

across the Pleistocene-Holocene transition. Results show that intensified use of small

game animals, especially rabbit, occurred much earlier than the end of the Pleistocene. In

addition, no discernible trend towards dietary diversification was found. Diets were

diverse during the entire Upper Paleolithic sequence in Iberia. It is argued that the

appearance of marine resource use at the end of the Pleistocene reflects changes in sea

level that have severely altered the archaeological record. The transport of marine resources

inland during the Early Upper Paleolithic shows that coastal resource use occurred much


For my Mom and Dad


It is not the critic who counts; not the man who points out
how the strong man stumbles, or where the doer of deeds could
have done them better. The credit belongs to the man who is
actually in the arena, whose face is marred by dust and sweat
and blood; who strives valiantly; who errs, and comes short
again and again; because there is not effort without error and
shortcoming; but who does actually strive to do the deeds; who
knows the great enthusiasms, the great devotions; who spends
himself in a worthy cause, who at the best knows in the end
the triumphs of high achievement and who at the worst. if he
fails, at least fails while daring greatly, so that his place shall
never be with those cold and timid souls who know neither
victory nor defeat.

-- Theodore Roosevelt, excerpt from his Speech at the

Sorbonne, Paris, France April 23, 1910

Many people inspired, assisted and facilitated this dissertation. It really began with

my first exposure to Paleolithic archaeology as an undergraduate at the University of

Arizona. The classes I took from Arthur Jelinek, John Olsen, Stanley Olsen, and Mary

Ellen Morbeck on Paleolithic archaeology and paleoanthropology sparked my interest in

this period of human prehistory. I never wavered too far off course despite numerous

temptations to explore other avenues.

Most of these temptations came to me in graduate school at the University of

Wisconsin-Madison. A wise man once said to me that choosing the right graduate school

was as important as choosing the right woman to marry. I have no doubt that I made the

correct decision. The influence of all of my faculty was most rewarding and I wish to

thank all of them: T. Douglas Price, James Stoltman, Mark Kenoyer, Henry Bunn, Gary

Feinman and Herb Maschner. They exposed me to the fascinating world of past human

behavior through numerous interesting and stimulating seminars and courses. I am

extremely grateful to them for passing on their knowledge and appreciation of the past.

Unfortunately, I never had the pleasure of taking courses from Sissel Schroeder and Jason

Yeager but I wish to thank them as well.

I would also like to thank the members of my dissertation committee for all of the

time they put in to reading and commenting on a previous draft. T. Douglas Price, Mark

Kenoyer, Henry Bunn, Sissel Schroeder and William Aylward helped make this a better

work with their kind, thoughtful and creative suggestions. They made what might have

been a stressful period truly pleasureable.


There are numerous institutions who helped make this work possible by giving me

opportunities and funding to continue research. The University of Wisconsin and its

Anthropology department provided travel grants that enabled me to go to Portugal to

explore my dissertation research. The Portuguese government supported the excavation

of Lapa do Picareiro through its grants to Nuno Bicho, without which most of my research

would not have been possible. Geochron Laboratories supported the radiocarbon dating

for the site, Lapa do Suão. The National Museum of Archaeology in Portugal kindly gave

permission to study materials and to take samples for further analyses. I thank all of these


A long time ago I begged on to the final year of a project on Upper Paleolithic

archaeology directed by Anthony Marks and João Zilhão in Portugal. It changed my life

immeasurably. Without that first opportunity to work in Portugal, none of this would

have happened and I am greatly appreciative. I thank Jeff Shokler for encouraging me to

contact Tony Marks to get started. While on that project, I met Nuno Bicho whom I have

had the greatest pleasure of working with ever since. The subsequent project he began

led to this dissertation. Many rewarding personal and professional contacts were

established during this last decade.

On a more personal level I wish to thank many people whom I worked, played and

became friends with. I am not really sure where to begin because there are so many to

thank. I would first like to acknowledge and thank all of my graduate school colleagues

and friends.

Jeff Shokler and I shared many conversations on Portugal, archaeology and food. I

thank him and Sherrie for feeding me wonderful food over the years.

Brian Hoffman and Matt Thomas have been great friends. I always enjoy our lunches

and hours of conversation.

I would like to give special thanks to my advisor, Doug Price for allowing me to grow

into the person I am. Without your support and guidance through the years I would

never have made it this far.

Nuno Bicho and I have worked and played together in Portugal for over 10 years. It

has been one of the most unimaginably wonderful experiences to explore Portugal with

him. I cannot possibly thank you enough and look forward to what I know will be many

years of friendship and further collaborative work.

I also thank Byran Hockett for what has turned out to be a beautiful professional and

personal relationship. Much of the intellectual quality of this dissertation is due to him.

Our collaborative work has been extremely rewarding. I value our work and friendship


I would also thank Maria João Valente for her generosity in allowing me to study part

of the Lapa do Suão collection. It has led to a warm and enduring friendship. Much of my

work in Portugal was made possible by Maria João and her in-laws, the Martins family.

In addition Antonio Faustino Carvalho and Francisco Almeida became good friends and

colleagues over the years. João Zilhão and Katia Araújo helped me in many ways and I

consider them friends and colleagues. I would also thank Paul Thacker for numerous

conversations and idea sharing about Portuguese archaeology.

It seems like Chris Fisher, Kiki Gilderhus, Mike Galaty, Caroline Funk and I have

been the best of friends forever. We have shared so many warm and fun experiences and

stimulating intellectual conversations. Since they have moved on I made fast friends with

David Meiggs, Susan Reslewic, Shawn Murray, Brad Chase and Kelly Knudson. I love all

of you and thank you from the bottom of my heart for being my friends and helping me

along the way.

Lastly, I would like to thank my entire family for their love and support. If not for

them I would never have done this. My sister Ann and my parents, Bob and Penny, gave

me everything I really need in life: love.


Table of Contents

Chapter 1: Introduction ............................................................................................................. 1

Part I: Theoretical frameworks for reconstructing subsistence and settlement

patterns, explaining subsistence change and past application on the Iberian
Peninsula .............................................................................................................................. 15

Chapter 2: Theoretical frameworks for hunter-gatherer subsistence and settlement

pattern studies ..................................................................................................................... 15
Site Catchment Analysis ......................................................................................................... 16
Forager/Collector Model ....................................................................................................... 17
Predictive models .................................................................................................................... 18
Optimal Foraging Models ...................................................................................................... 24
Diet Breadth .............................................................................................................................. 25
Patch Choice ............................................................................................................................. 27
Central Place ............................................................................................................................. 28
Discussion ................................................................................................................................. 30
Nutritional ecology and human dietary choice .................................................................. 33
2.1 Explanations for subsistence change ................................................................................ 38

Chapter 3: A regional perspective on Upper Paleolithic Iberia: a comparison of

Cantabria, Mediterranean Spain and Portugal ............................................................. 46
3.1 Archaeological applications of economic models in Spain ........................................... 46
Cantabria ................................................................................................................................... 47
Mediterranean Spain ............................................................................................................... 52
3.2 Central Portugal ................................................................................................................... 62
Subsistence ................................................................................................................................ 69
Settlement ................................................................................................................................. 72

Part II: Prehistoric diet and subsistence................................................................................76

Chapter 4: A Western Mediterranean perspective on Upper Paleolithic plant

consumption ........................................................................................................................ 81
4.1 Plant use by prehistoric Mediterranean hunter-gatherers ............................................ 81
4.2 Plant resources in Iberia ..................................................................................................... 85
Present-day climate and vegetation in Portugal ................................................................. 85
4.3 Late Pleistocene/ Early Holocene Portugal ................................................................... 102
Climate change and paleoenvironment ............................................................................. 103
4.4 Modeling economic and nutritional utility of plants ................................................... 124
Pine nuts .................................................................................................................................. 129
Acorns ..................................................................................................................................... 132
Nutritional utility .................................................................................................................. 138
4.5 Discussion ........................................................................................................................... 141

Chapter 5: Upper Paleolithic faunal exploitation in central Portugal .......................... 146

5.1 History of investigation .................................................................................................... 146
Pleistocene faunas .................................................................................................................. 145
5.2 Prey behavioral ecology.................................................................................................... 150
5.3 Lapa do Picareiro ............................................................................................................... 156
Faunal Remains ...................................................................................................................... 164
Taphonomy............................................................................................................................. 172
Skeletal element representation and butchery patterns ................................................... 183
Discussion ............................................................................................................................... 210
5.4 Lapa do Suão.......................................................................................................................213
Faunal analysis for Lapa do Suão.........................................................................................219
Rabbits ..................................................................................................................................... 222
Avifauna.................................................................................................................................. 236
Aquatic fauna......................................................................................................................... 239
Discussion............................................................................................................................... 239
5.5 Additional fauna-bearing Final Upper Paleolithic and Epipaleolithic sites in
Portuguese Estremadura ........................................................................................................ 241
Gruta do Caldeirão ................................................................................................................ 242
Bocas, Casal Papagaio, Pena da Mira ................................................................................. 248
Lapa dos Coelhos ................................................................................................................... 253
Buraca Grande & Buraca Escura ......................................................................................... 253
5.6 Discussion ........................................................................................................................... 254
5.7 The broader regional context ........................................................................................... 258

Chapter 6: Towards an understanding of Late Pleistocene/ Early Holocene

subsistence and settlement in central Portugal .......................................................... 272
Summary ................................................................................................................................... 293

References Cited ...................................................................................................................... 299


Chapter 1: Introduction

This study utilizes archaeological data to test the efficacy of the Broad Spectrum

Revolution (BSR) model in central Portugal during the Late Pleistocene and Early Holocene.

It explores the relationships between humans and their surrounding environment in an

evolutionary ecological framework that encompasses traditional paleoeconomic models,

optimal foraging theory and nutritional ecology. The Late Upper Paleolithic and

Epipaleolithic diet, subsistence and settlement patterns of central Portugal are examined

through paleoenvironmental data and detailed studies of archaeofaunal assemblages

through taphonomic lenses. The goal of this dissertation is to show that the main

components of the Broad Spectrum Revolution model, resource intensification and

diversification, did not suddenly appear at the beginning of the Holocene, but that they

have a much greater time depth. It is argued here that dietary diversity is part of our

evolutionary heritage as omnivorous primates and shifts between generalized and

specialized diets reflect local climatic and environmental conditions, not a directional trend

in human adaptation.

Archaeologists have long characterized the Pleistocene/Holocene transition as a

boundary across which human societies underwent dramatic changes that ultimately led

to the transition to agriculture. These include the intensification and diversification of the

human subsistence base and increased sedentism, seen by greater logistical mobility, larger

residential site size and the appearance of cemeteries. The most significant aspects of pre-

agriculture human subsistence diversification are the worldwide increase in the exploitation

of small game and aquatic resources and more intensive use of plant foods, especially

seeds and nuts (Binford 1968; Flannery 1969; Hayden 1981; Bailey and Parkington 1988;

Kuhn and Stiner 2001). Flannery (1969) characterized this global trend as a ‘Broad Spectrum

Revolution’ in human diet. Ultimately, this dietary shift is thought to result from human

population pressure on resources. As population increased in the Late Pleistocene, people

were forced to incorporate new food items into their diet that were previously ignored.

Increasingly, archaeologists are pushing the exploitation of small game, aquatic resources

and plants further back in time (Erlandson 2001; Stiner 2000). Recently, Stiner et al. (2000)

have modified the Broad Spectrum Model to account for the discrepancy between the

archaeological evidence and the expectations of the original model. An alternative

explanation to the nature and timing of small game, aquatic and plant resource exploitation

has been offered by Hockett and Haws (2002, in press). In this approach, dietary diversity

is seen as a hallmark of human adaptation not the result of progressive trends due to

gradual population increase. Because these developments occurred on a global scale,

information from a wide range of geographic and environmental contexts is required in

order to better understand them (Straus 1996). The focus here is on the Iberian Peninsula

(Figure 1.1).

On the Iberian Peninsula, the overwhelming majority of the research on subsistence/

settlement patterns during the Late Pleistocene/Early Holocene comes from Cantabria

and Mediterranean Spain (Aura Tortosa & Pérez Ripoll 1995; Bailey 1983; Bailey and

Davidson 1983; Clark 1983a, 1983b; Conkey 1980; Moure-Romanillo 1995; Straus 1986,1987;

Straus and Clark 1986; Villaverde and Martinez Valle 1995; Villaverde et al. 1998). This

R Ebro
R Douro

Atlantic Ocean

R Tejo València
R Júcar


Mediterranean Sea




Figure : Map of the Iberian Peninsula


research was facilitated by good faunal and other organic preservation in caves and

rockshelters. Diachronic changes in subsistence and settlement patterns are characterized

by gradual diversification and intensification of the subsistence base from the Solutrean

through the Mesolithic (Clark 1987; Clark and Straus 1986; Clark and Yi 1983; Straus 1992).

Clark and Straus (1986) explained subsistence change as a result of demographic expansion

and resource stress, largely based on the faunal record of red deer specialization and the

increase in shellfish use at La Riera cave, thus supporting the ‘population pressure’ theory

and the ‘Broad Spectrum Revolution’ model for the Late Pleistocene/Early Holocene

(Binford 1968; Clark and Straus 1986; Cohen 1977; Earle 1980; Hassan 1981; Neeley and

Clark 1993). In Mediterranean Spain, researchers have observed a similar Late Pleistocene

trend of intensified rabbit, red deer and ibex exploitation (Aura and Villaverde 1995; Aura

and Pérez Ripoll 1992, 1995; Aura et al. 1998, 2002; Villaverde and Martínez Valle 1992,

1995; Villaverde et al. 1997, 1998). In this region however, there is evidence of plant and

marine resources in archaeological sites dated to the Middle Paleolithic in Gibraltar

(Finlayson et al. 2000). Coastal and plant resources are also apparent in the Upper Paleolithic

levels of the Cueva de Nerja. This suggests the Broad Spectrum Revolution model may

not fit the data from Mediterranean Spain.

One of the areas of visible post-Pleistocene shifts towards coastal resource use is

the central region of Portugal (Binford 1968; Clark 1952). Large shell middens dated to

the Early Holocene near the Tejo estuary have been known for almost 150 years. Their

appearance is thought to result from human dietary and settlement shifts coinciding with

the global pattern described as the Broad Spectrum Revolution. However, Portugal has

received comparatively less attention, yet remains the most appealing sub-region on the

peninsula to study the long-term changes outlined above. Initial investigation into the

Paleolithic of Portuguese Estremadura began in the 19th century by the geologists, Carlos

Ribeiro and Joaquim Filipe Nery Delgado (Bicho 1993; Zilhão 1993). Their work, and that

of the Serviços Geologícos, resulted in the discovery and excavation of many Upper

Paleolithic sites (Zilhão 1993). Numerous caves were excavated around the turn of the

century by Vieira Natividade and others from the Serviços Geológicos in the area between

the coast and Serra de Candeeiros. The region was then largely ignored until the 1930s

and 1940s, when Manuel Heleno excavated sites near Cambelas (Torres Vedras) and in

the Rio Maior valley. Perhaps the greatest impact on Portuguese Paleolithic archaeology

came from the work of George Zbyszewski and Henri Breuil. Using the fossile-directeur

approach to typology and chronology, Breuil classified the Portuguese Upper Paleolithic

in the same terms as the classic French core area. Despite differences between the two

regions, the sequence of Aurignacian (33-25,000 bp), Gravettian (25-22,000 bp), Solutrean

(22-17,000 bp), and Magdalenian (17-10,000 bp) is used to the present day to organize the

Upper Paleolithic. Epipaleolithic (10-8,000 bp) has been added to this repertoire to classify

sites dated to the Early Holocene which are still characteristic of the Final Upper Paleolithic

(Bicho 1996; but see Vierra 1995 for a different scheme). The regional cultural chronology

is provided in Table 1.1.

Interest in the Late Upper Paleolithic and Mesolithic waned until the 1980s, when

research begun by João Zilhão, José Morais Arnaud, and David Lubell contributed

significant information on the prehistory of central Portugal (Arnaud, 1993; Bicho, 1993b;
Table : Cultural divisions for Late Pleistocene/Early Holocene southwest
Southwest France Northern Spain Mediterranean Spain Portugal

Mesolithic Asturian/Geometric Geometric Epipaleolithic Mesolithic

(Sauveterrian) Epipaleolithic 
 bp  bp
 bp  bp
Azilian Azilian Microlaminar Epipaleolithic Final Magdalenian/
 bp  bp   bp Epipaleolithic
Upper Magdalenian Upper Magdalenian Upper Magdalenian Late Magdalenian
  bp   bp   bp   bp
Lower Middle Lower Cantabrian SolutreoGravettian Early Magdalenian
Magdalenian Magdalenian Magdalenian   
Solutrean Solutrean Solutrean Solutrean
 bp   bp

Adapted and modified from Straus ()


Lubell and Jackes, 1985; Marks et al. 1994; Zilhão, 1992,1993). Most recently, research has

focused on chronology, lithic typology and technology (Bicho, 1992; Marks et al., 1994;

Zilhão, 1990, 1991, 1995). This work resulted from a collaborative project between Anthony

Marks and João Zilhão, funded by the National Science Foundation. The Upper Paleolithic

of Portuguese Estremadura Project surveyed and excavated many Upper Paleolithic

(33,000-10,000 years ago) sites near the research area (Marks et al 1994).

Over the last fifteen years new syntheses of old information and results of new

excavations have been used to develop explanatory models for Late Upper Paleolithic,

Epipaleolithic and Mesolithic human adaptation in central Portugal (Arnaud 1986, 1989,

1993; Aura et al. 1998; Bicho 1993, 1994, 1996, 1997; González Morales and Arnaud 1990;

Zilhão 1990, 1992, 1995; 1997). Several recent studies focused on hunter-gatherer mobility

and movement by studying lithic technology, raw material procurement and limited

information on subsistence (Bicho 1992, 1993, 1994, 2000; Shokler 1995; Thacker 1996, 2000;

Zilhão 1990, 1992, 1995). Vierra (1995) attempted to link changes in lithic technology to

subsistence strategies. Understanding Late Pleistocene/Early Holocene subsistence and

settlement patterns requires consideration of subsistence patterns together with raw

material procurement, lithic assemblage variability, and site location. Each of the models

outlined suffers from the fact that they lack crucial data on subsistence and coastal site


Preliminary faunal analyses from Lapa do Picareiro and impressions from other

sites resulted in a model of intensified rabbit and red deer procurement and a broadening

of the diet to include aquatic resources and possibly plants at the end of the Upper

Paleolithic in central Portugal (Bicho and Haws 1996). However, subsequent work suggests

that these resources were part of the diet for the entire Upper Paleolithic and may date to

the Middle Paleolithic (Hockett and Haws 2002). In fact, little is known about human

subsistence during the Late Pleistocene and Early Holocene in Portugal. The visible changes

in the archaeological record have been interpreted without the benefit of careful taphonomic

and behavioral studies of the faunal assemblages. Only recently have systematic and

taphonomic analyses of Upper Paleolithic faunal assemblages been conducted (Aubry et

al. 2002; Bicho et al. 2000; Davis 2002; Haws 1998; Hockett and Bicho 2000; Valente 2000).

These studies, and the analyses presented in this dissertation, are used to reevaluate the

validity of the proposed models for the region centered on the larger question of Late

Pleistocene/Early Holocene subsistence change. Some new interpretations using a

nutritional ecology approach have already been proposed (Bicho et al. 2002; Hockett and

Haws 2002a, 2002b, in press).

Portuguese Estremadura contains numerous Upper Paleolithic and Epipaleolithic

sites indicating it was an important focus of prehistoric human occupation in Iberia. Other

areas were probably occupied but systematic surveys to discover sites outside Estremadura

have only begun in the last five years. Recent models of Late Pleistocene/Early Holocene

subsistence and settlement patterns in central Portugal are based on the BSR model, namely,

that resource intensification and diversification took place because of population pressure

on resources. If resource intensification and diversification occurred, it should be visible

in the patterning of food refuse in archaeological sites. To test the general model, this

study uses faunal analyses of two Late Upper Paleolithic sites, Lapa do Picareiro and

Table  : Components of the BSR model and its manifestation in Iberian prehistory followed by
a contrasting interpretation using a nutritional ecology approach

Broad Spectrum Revolution model Broadening of the diet at the PleistoceneHolocene

transition or later
Caused by increased human population due to inherent
growth trends in modern humans
Archaeological manifestation is the appearance of small
game (rodents and birds) aquatic resources (shellfish)
and plant (nuts and seeds)

Stiner modifications Introduced new version of the model to account for the
early appearance of small game and shellfish
Periods of low population characterized by the use of
slowmoving ‘sessile’ resources like tortoises and shellfish
Size diminution in the slowmoving animals indicates
overharvesting due to population pressure
Diet breadth model accurately predicts a shift from
tortoises and shellfish to ‘quick’ hard to catch lagomorphs
and birds

Appearance of fastmoving resources signals population


Iberia Resource intensification diversification and specialization

apparent as early as the Solutrean in Cantabria
Evidence of shellfish overharvesting seen by size
Increased number of sites in the Solutrean period suggests
increased regional population due to influx of people from
northern Europe
Mediterranean Spain has evidence for resource
intensification (rabbit) diversification (shellfish and
birds)and specialization (rabbit red deer and ibex)
beginning in the Magdalenian

PleistoceneHolocene transition in Portugal is

characterized by resource intensification (rabbit red
deer) diversification (shellfish) and specialization (rabbit
red deer)
Population increase seen by increased number of sites per
millennium and expansion into previously unoccupied
Population pressure responsible for dietary shifts

Nutritional ecology model Dietary diversity in the Paleolithic predates evidence for
population increase It is an inherent feature of human
dietary evolution
Human populations increase because balanced essential
nutrient intake leads to lower infant mortality rates and
better reproductive success
Population increase is not a constant feature of human
Dietary shifts are responses to changing resource
availability and the need to maintain balanced nutrient
Dietary shifts in the Late Pleistocene correspond to
changing environments
Increases in the numbers of sites in the Late Pleistocene
are a function of rising sea level and site visibility

Lapa do Suão, paleoenvironmental data from deep sea cores, pollen diagrams, charcoal

analyses and inferences about human behavior derived from evolutionary ecology. The

faunal assemblages from the two sites form the best source of data because the ones from

the other previously excavated sites were either partially recovered or the collections do

not survive. One site, Gruta do Caldeirão, has recently been analyzed and these results

are discussed along with others under excavation that have not been fully reported. Because

the overall sample in Portuguese Estremadura is small, a regional perspective is taken in

order to better understand variability in human dietary choice and subsistence behavior

on the Iberian Peninsula. It will be shown that while some of the Estremaduran caves

contain well-preserved faunal remains, they are not fully representative of the full range

of subsistence behavior. This has important implications for each of the models for Portugal

and the larger issues of intensification, diversification and specialization incorporated in

the BSR model.

Table 1.2 shows some expectations of the original Broad Spectrum Revolution model

and recent modifications along with the archaeological manifestations. It also lists the

conclusions based on previous application of the model to the Iberian Peninsula. Lastly,

alternative expectations are presented from a nutritional ecology perspective. Whatever

the explanation for subsistence change in prehistory, it is clear that detailed taphonomic

studies must be done before higher level explanatory models can be critically evaluated.

Less than systematic recovery and curation of faunal and floral remains from archaeological

sites can lead to erroneous conclusions about human behavior. Unfortunately, many of

the studies designed to test ideas about past human behavior are based on such faulty


The organization of this dissertation is two fold. The first part concerns theoretical

and methodological approaches to the study of hunter-gatherers on the Iberian Peninsula.

The second part is an exploration of the archaeological record of Late Upper Paleolithic

and Epipaleolithic central Portugal.

Chapter 2 traces the development of theoretical approaches to studying prehistoric

hunter-gatherers. It begins with the work of Grahame Clark and the rise of the Cambridge

paleoeconomy school from the 1950s to the 1980s. Site catchment analysis is specifically

discussed because of it application to Iberia. Afterwards, the influential work of Lewis

Binford on hunter-gatherer subsistence and settlement patterns is discussed. Following

this is a detailed review of Jochim’s (1976) explanatory model for hunter-gatherer

subsistence and settlement patterns, which illustrates the level of analytical intricacy that

is necessary to fully grasp hunter-gatherer decision-making. This leads into a general

introduction to the models collectively known as optimal foraging theory under the larger

umbrella of evolutionary ecology. Included in the discussion is a new approach called

nutritional ecology recently offered by Hockett and Haws (2002, in press). This approach

is also appropriately placed within evolutionary ecology. Though discussed later in more

detail, this model was proposed to account for non-energetic nutritional concerns in human

dietary choice that were raised by linear programming proponents in the late 1970s and

early 1980s.

The purpose of this discussion is to provide a framework for the analyses and their

interpretation, not to expicitly test each model. Many of these are untestable using the

archaeological record but have been useful heuristic devices to make inferences about

past human behavior.

Explanations for diachronic subsistence change and the merits of each are discussed

in the second part of chapter 2. The discussion centers on the Broad Spectrum Revolution

model and the nutritional ecology approach. The BSR is essentially the same as the diet

breadth model in foraging theory. Both are used by archaeologists to understand human

dietary choices based on the capture of energy from the environment. The nutritional

ecology approach expands the currency to include essential nutrients required by the

human body to maintain health and ensure reproductive success.

Chapter 3 illustrates the application of the theoretical and methodological

approaches to the Late Pleistocene and Early Holocene of the Iberian Peninsula. It begins

with a brief review of the models of subsistence and settlement patterns developed by

Lawrence Straus, Geoffrey Clark and Spanish colleagues. A review of Mediterranean

Spain includes a review of the application of site catchment analyses by Geoff Bailey and

Iain Davidson and their impact on prehistoric hunter-gatherer research by Spanish

archaeologists. These studies were instrumental in determining that medium ungulate

exploitation was highly specialized during the Late Upper Paleolithic. In both regions

Anglo-American archaeological thought has had a powerful influence. Many of the ideas

concerning trends in Paleolithic societies stem from the theoretical approaches outlined in

chapter 2. These are evident from the works of Valentin Villaverde Bonilla, Rafael Martínez

Valle, Emili Aura Tortosa and Manuel Pérez Ripoll. They have all done extensive research

into the long term trends in subsistence and settlement strategies throughout the Paleolithic

in the Spanish Mediterranean Region.

The last part of this chapter introduces the models developed by archaeologists to

explain Upper Paleolithic hunter-gatherer subsistence and settlement pattern in central

Portugal. The most comprehensive models were proposed by João Zilhão and Nuno Bicho

in the last decade. Their approaches reflect differences in European (mainly French) and

American perspectives on the past. Both note the intensification and specialization of

resource use along with diversification at the end of the Pleistocene. Their models are

presented and discussed.

Part II has a short introduction to archaeological and ethnographic perceptions of

hunter-gatherer diets through time. This section includes chapters on plant and animal

exploitation in central Portugal. Chapter 4 begins with a regional survey of the evidence

for plant exploitation from Pleistocene and Early Holocene sites in the Mediterranean.

This is followed by a discussion of the present-day environment in Iberia and characteristics

of some edible wild plants that were economically important in the past. Understanding

the availability of plant resources for Upper Paleolithic people in central Portugal requires

detailed paleoenvironmental reconstructions. Data from deep sea cores, pollen cores,

charcoal analyses, microfaunal and macrofaunal inventories are presented in order to

reconstruct Late Pleistocene climate and environments in Iberia. Afterwards, expectations

from ethnographic observations and foraging models are used to interpret the available

data from Upper Paleolithic archaeological contexts.


Chapter 5 focuses on the exploitation of terrestrial animal resources in central

Portugal during the Late Upper Paleolithic. It begins with a brief introduction to the

previous research on animal bones from Upper Paleolithic sites in Portugal. A brief

discussion of the behavioral ecology of prey selected by human hunters is intended to

help understand prehistoric decision making and assemblage composition in each site.

Faunal analyses of two sites, Lapa do Picareiro and Lapa do Suão, are discussed in detail.

These are the best sources of information for Magdalenian subsistence in all of Portugal.

The analysis focuses on the taphonomy and skeletal element representation of key taxa in

order to explain the assemblage composition. For each site the role of carnivores and

potential effects of other agents of density-mediated bone loss are discussed before

interpretations concerning human treatment of animal carcasses are made. After these

assemblages are discussed, they are compared to other sites in Portuguese Estremadura.

The Caldeirão Cave assemblage is given special attention. The last section of chapter 5 is

a discussion of the subsistence strategies observed in central Portugal and comparison

with those from Mediterranean Spain.

Chapter 6 concludes the dissertation by applying the subsistence and settlement

data from Portugal against the Broad Spectrum Revolution model and others presented

in detail in chapter 3. A hypothetical model of Late Pleistocene and Early Holocene

subsistence in central Portugal is presented. Finally, some directions for future research

are outlined.

Part I: Theoretical frameworks for reconstructing subsistence and settlement

patterns, explaining subsistence change and past application on the Iberian


Chapter 2: Theoretical frameworks for hunter-gatherer subsistence and settlement

pattern studies

One of the most fertile domains of research on Paleolithic hunter-gatherers is found

in human-land relationships, such as the construction of models of subsistence/ settlement

patterns (Bettinger 1991; Binford 1980, 1982; Gamble 1986; Jochim 1976, 1998; Kelly 1995;

White 1987). Paleolithic hunter-gatherers are investigated primarily through subsistence

(faunal and floral remains), technological organization and raw material procurement

(stone tools), and paleoenvironmental reconstruction (geological, palynological, and

climatological studies). Subsistence and settlement pattern studies generally involve

economic considerations based on the capture of energy from the surrounding

environment. Most models incorporate modified versions of site catchment analysis (Vita-

Finzi and Higgs 1970), the collectors vs. foragers model (Binford 1980), or optimal foraging

theory (Winterhalder and Smith 1981). Although the primary focus for study is the

economy of hunter-gatherers, a more complete understanding of past behavior should

include the social and ideological realms. However, these issues are difficult to address

due to the lack of material preservation (but see Gamble 1999 for an alternative view).

The following sections serve as a review of economic approaches to studying prehistoric


subsistence and settlement patterns. These approaches have been used to interpret the

Upper Paleolithic archaeology of the Iberian Peninsula and to explain diachronic trends

during this period.

Site Catchment Analysis

Beginning with the seminal paper by Vita Finzi and Higgs (1970), site catchment

analysis became a primary method used by prehistorians to reconstruct (or construct)

paleoeconomies. The method was designed to “study the relationships between technology

and those natural resources lying within economic range of individual sites” (Vita Finzi

and Higgs 1970: 5). It assumes that humans will exploit “those resources in its territory

that are economic ... and that are within reach of the available technology” (Vita Finzi and

Higgs 1970: 2). Territories are considered to be limited by space and time in that there is

an average threshold in the distance and length of time people will travel from home

bases in their daily quest for necessary resources. In hunter-gatherer societies it has been

observed ethnographically that people tend to stay within 10 km or 2 hours walking time

of their home base. Bailey (1983a) has made an important, if somewhat confusing,

distinction between site catchment analyses and site exploitation territory analyses. The

former represents an “empirical statement of the site exploitation territory, while the site

exploitation territory represents a hypothetical assessment of the economic catchment”

(p.61). According to Bailey and Davidson (1983), what is commonly thought of by most

archaeologists as ‘site catchment analysis’ is called ‘site territorial analysis.’ In other words,

a site territorial analysis is an examination of the potentially available resources within


the vicinity of the site. The site catchment is then argued by Bailey and Davidson to be the

actual area from which resources found in an archaeological site are procured.

Bailey and Davidson (1983) state six objectives of site catchment analysis. These

1) to define the area habitually used by the occupants of a site for their daily
2) to trace to their points of origin in the surrounding landscape materials and resources
whose archaeological remains are present on-site;
3) to reconstruct the microenvironments around a site as a clue to variations in the
environmental data present on-site;
4) to reconstruct the food resources potentially available to the occupants of a site
and hence, by further inference, the subsistence economy actually practiced by them;
5) to reconstruct the function of sites (as home-bases, temporary camps, etc.);
6) to reconstruct the social and economic relationships between sites as members of
regional settlement systems (p.88).

Forager/Collector Model

Binford (1982) later modified the simplistic site catchment model. He wrote:
It is unrealistic to view the potential zonation around a residential camp as simply a
series of concentric circles where the use which is made of each area is exclusively
conditioned by the transport and labor costs of exploiting resources at differing
distances from a locus of consumption (see Jochim 1976: 51-56). The situation is
more realistically visualized as a residential camp at the hub of a foraging radius
and a logistical radius (Binford 1982: 8).

In Binford’s scheme, there are two end points along a continuum. Collectors employ

a settlement strategy of low residential mobility with a high degree of logistical mobility.

In this case, people live in relatively large residential base camps and foraging would take

place in the immediate zone around the camp, usually within a 2-hour walk. Task groups

make logistical forays into areas at greater distances requiring overnight or longer stays

away from the main camp (Binford 1982). Several site types should be archaeologically

visible including butchery/processing camps, stations, caches, etc. Conversely, foragers


tend to live in smaller groups and move their residence often. This usually occurs before

resources in the immediate vicinity of a camp are depleted. Ultimately, length of stay in a

residential camp will be dependent upon the availability of resources within the logistical

radius of the site. The two general site types created are the residential camp and location.

The former is where the majority of daily tasks are undertaken and the latter are specific

points on the landscape where foragers go to obtain certain resources and return within a

few hours. In this case overnight stays are considered rare. Generally, resource distribution

conditions whether groups adhere to the forager or collector pattern. Foragers usually

are found in environments with even resource distribution, while collectors typically inhabit

patchy environments, although these are not mutually exclusive divisions (Binford 1980).

Binford (1978) proposed methods for identifying settlement types based largely on

faunal remains. His ethnographic work on the Nunamuit and subsequent experimental

work resulted in utility indices for understanding observed skeletal element patterning.

The Modified General Utility Index (MGUI) is an economic measure of body part value.

Butchering strategies are conditioned by supply and demand. In cases of high supply/

low demand the higher utility body parts will be consumed or transported to a residential

camp resulting in an assemblage dominated by low utility parts. This is referred to as a

gourmet curve. When supply is low and demand high, a more intensive utilization of

body parts results in a bulk curve (Binford 1978; Bettinger 1991).

Predictive models

Predictive models of hunter-gatherer behavior have been used in the past three decades

to understand subsistence and settlement strategies. Jochim (1976) borrowed concepts of


efficiency and risk from economic anthropology and built a decision-making model using

ethnographic observations to predict prehistoric land use. His model involved three main

assumptions: (1) Resource choice structures hunter-gatherer subsistence and settlement

patterns. Choice requires decisions made to allocate time and energy for resource

acquisition and utilization as well as group size and spacing; (2) choice usually is based

on long range, goal-oriented decisions and rarely is opportunistic (immediate); (3) people

are rational decision-makers. Making rational decisions requires knowledge of risks

involved and possible outcomes. Jochim places hunter-gatherers in a state of ‘partial

uncertainty’ since the exact probability of the outcomes of economic choices can be known

only from previous experience (p.5). The main goals in a hunter-gatherer economy are

minimizing effort and risk. With these principles in mind, the organization of hunter-

gatherer subsistence and settlements patterns are placed in an ecological framework using

systems theory.

The decisions made by a hunter-gatherer population address essential problems that

need to be solved. These include: the resources used and the amount of each needed, the

timing and location of resource use, and group size and labor division. Three main

subsystems include resource use schedule, site placement and demographic arrangement.

The priority of the system lies with the resource use schedule, which requires the

identification of specific goals, resources exploited, environment, measures of performance

and management considerations. The two main goals are providing enough calories to

sustain the population and exploiting the most reliable resources, those which provide

the highest return and least risk. Additionally, resources that can be procured through

the least amount of work are preferred. Often this is accomplished by exploiting easily

captured resources, timing the use to coincide with peak abundance of the resource, and

reducing the costs of travel, mainly just distance, but this point will be discussed later in

more detail. Effort placed into food procurement is also conditioned by the need for

population aggregation for mate exchange, food sharing, cooperative hunting, trade and

rituals. Of secondary importance are the desires for taste, variety, prestige and sex role

differentiation. For Jochim, taste is largely based on fat content of animals. Speth and

Spielmann (1983) consider fat a dietary necessity and this need may condition subsistence

and settlement strategies during stressful times of the year, mainly the late Winter and

early Spring.

The resource use schedule is largely determined by the behavior of the resource (Jochim

1976: 22-33). The key attributes are weight (w), density (d), aggregation size (a), mobility

(m), fat content (f) and additional nonfood characteristics. Of these, weight and density

are the two attributes considered in attaining the nutritional goals. The population density

and mobility of a resource determine the search and capture probability enabling risk

management decisions. Aggregation is possible where high yield resources are

concentrated. This in turn is conditioned by the weight and nonfood yield, resource

aggregation size and mobility or distance to intercept.

In Jochim’s model, environmental factors such as climate, geography and seasonality

cause variation in the resource use schedule. Seasonality is the most important because it

“induces a patterned variation of the resource attributes” (Jochim 1976: 24). Plants are

affected by the growing season. The weight and fat content of animals change with seasonal

availability of their food, and reproductive cycles lead to variation in group size and

mobility of animal populations. Nonfood attributes are also affected by seasonal cycles.

Jochim then provides a mathematical means of estimating the performance of resource

attributes in attaining the goals of the decision-making process. Each attribute is considered


Secure food and nonfood income wnd/m

Population aggregation at minimum cost wna/m

Taste f


Prestige wnfm/d

Sex role differentiation

He then scores each resource in order to determine their proportional use.

resource use %= resource score/sum of scores x 100
total resource use %= sum of 2 resource use %’s/2

Again, reliability and least cost are the two most important goals of resource use. The

common denominator of both is the resource mobility, either in absolute terms of distance/

time or the predictability in locating a resource as per its spatial behavior. In other words,

search and pursuit/capture costs. Seasonal changes in weight, aggregation and mobility

are corrected for by calculating resource use percentages in monthly time intervals. A

fairly good fit between the predicted resource utilization based on these measures calculated

from game in northern Ontario and actual resource use percentages by Round Lake Ojibwa

showed the potential utility of Jochim’s model.

The second subsystem of the model concerns settlement location (Jochim 1976: 47-

63). The primary goals include proximity of economic resources, shelter, and view.

Generally, economic resources are food, although in arid or cold environments, water and

fuel may be of equal or greater importance. Jochim uses a gravity model to predict

settlement location based on ethnographically observed organizational structure. The

formula I=M1M2/R2 measures the interaction (I) of two populations (M) and the distance

(R) between them. Then I= kp or the proportional use of a resource. The mass of a

population is a constant (K) whereas the resource mass equals wna. Therefore, kp= Kwna/

R2. Rearranged to solve for distance and dropping the constants gives R2= wna/p. The

location of a settlement is determined by the resource attributes and its dietary proportion.

A hierarchical resource evaluation based on mobility results in a settlement pattern of site

location near less mobile, denser (more abundant) and less clustered resources. Preferred

site location is near low risk, low prestige food resources such as plants, small game or

fish with access to higher risk, higher prestige resources like big game. In Binford’s terms,

the more reliable resources would be found within the foraging radius of a base camp,

while the higher risk large game are within the logistical radius.

Settlement location is also chosen with regard to caloric requirements of a population.

The catchment or exploitation radius needed to fulfill the caloric requirements for a given

length of stay is measured by resource potential. Jochim considers the harvesting efficiency,

dietary proportion and kilocalories per km2 provided for each resource to arrive at figures

slightly larger than those estimated by Vita-Finzi and Higgs (1970). However, the prediction

of a winter exploitation radius of 16 km2 based on moose fit the observed Ojibwa pattern.

Again, the seasonal availability of resources influences the decision to settle a particular

region. The pull of a resource in other areas and exhaustion of local resources will affect

the occupational duration of an area. This also is dependent on human and resource

population density plus harvest efficiency, which, in turn, is dependent on population

and technology.

The demographic arrangement of a settlement system is dependent on the resource

use schedule (Jochim 1976: 65-79). Population density and spatial distribution are not

determined by the environment but rather the human decision making goals of food

provision, minimizing effort and risk, reproduction and social interaction. The fulfillment

of the goals requires the calculation of carrying capacity, which is derived from the biomass

of each resource in relation to dietary proportion, and their abundance and spatial

distribution. Biomass is measured by the kilocalories per 100 km2 for each resource.

Dividing by its proportion in the diet gives the carrying capacity. The maximum

supportable population is calculated by dividing the carrying capacity by the yearly caloric

requirements for one person based on an average of 2,000 kcal/person/day. The lowest

maximum supportable population figure is the highest possible population density. This

part of the model assumes 100% harvest efficiency or cull rate which is unsustainable and

must be corrected for. A lower efficiency of 27% for moose (the lowest figure) fits the

predicted and observed Ojibwa population density.

Population aggregation and dispersal will depend on the resource use schedule as

well. In order to minimize costs, group size is determined by the abundance of resources

at any given time. Adding resource density to the measure for low-cost population

aggregation (wnad/m) for each resource allows the estimation of group size. The sum of

the values for each resource per season enables the prediction of hunter-gatherer


Because of its comprehensive coverage of hunter-gatherer subsistence, settlement and

demography, Jochim’s model was applied to other archaeological cases. Although, the

basic concepts of efficiency (least effort) and reliability (risk minimization) remain central

to most models of prehistoric hunter-gatherer behavior, explicit use of his model has waned

in favor of other approaches to hunter-gatherer subsistence such as optimal foraging theory.

Optimal Foraging Models

In the last 20 years, many archaeologists studying hunter-gatherer subsistence have

adopted an evolutionary ecology approach. Evolutionary ecology is defined by

Winterhalder and Smith (1992) as, “the application of natural selection theory to the study

of adaptation and biological design in an ecological setting” (p.5). The most commonly

utilized set of models applied to archaeology are optimal foraging models borrowed from

ecology (Broughton and O’Connell 1999; Winterhalder 1981). Like Jochim’s, these models

have their roots in economic theory (MacArthur and Pianka 1966). The main assumption

of optimal foraging theory is that humans adapt to environments to harvest resources

with maximum efficiency. The goal of behavior is to achieve maximum fitness. Adaptation

is measured in net energy capture, while success is determined by the net acquisition rate

(Winterhalder 1981). Though energy is the most common currency, others such as

nutritional quality, in particular fat content, can also be used although they almost never

are (Winterhalder 1981; Speth & Spielmann 1983; Jochim 1976). The three models of interest

here are the Diet Breadth Model, Patch Choice Model and Central Place Foraging Model.

Diet Breadth

Hunter-gatherers living in a specific environment must decide which food resources

to pursue and which to ignore. The Diet Breadth model predicts whether or not a resource

should be pursued after encounter. In the model, procurement costs of food resources in

a given environment are divided into search costs (time in locating the resource) and

handling costs (time in pursuit, capture and processing) (MacArthur and Pianka 1966;

Winterhalder 1981). Items are included in the diet as long as the handling costs are

compensated by decreased search costs. The main difference between the two is that all

prey are searched for, but only the higher ranked ones are pursued. Search costs are

based on the resource density and mobility. Handling costs are also affected by processing

technology. High search costs generally result in broad diet breadth whereas high pursuit

costs result in narrow diets. Search costs are often conditioned by the “patchiness” of the

environment so that foragers living in “fine-grained” environments typically have broad

or generalized diets while those in “coarse-grained” ones have narrow or specialized diets

(MacArthur and Pianka 1966; Winterhalder 1981).

The model assumes that the primary goal of foraging is the maximization of energy-

return rates (Kelly 1995). Resources are then ranked according to post-encounter return

rates, based on the energy (kilocalories in a single unit of resource multiplied by the amount

acquired) per unit time after encounter, usually kcal/hr. Despite long search time, large

game are usually the highest ranked resource because the caloric yield is great enough to

significantly offset handling costs. Small game are often more abundant and thus have

lower search costs, but pursuit costs are usually higher because more effort goes into

capturing and handling them. Also, their small package size results in a low post-encounter

return rate requiring a higher harvest rate to make it the worth the effort. Plant foods,

though abundant and often very reliable, rank even lower in post-encounter return due to

an even lower caloric yield per item, thus need to harvest and process large quantities

which increases the pursuit costs. Therefore, the abundance of a resource does not

determine its inclusion in the diet. Extremely abundant, low-ranked resources may be

ignored as long as high-ranked resources are sufficiently abundant. Lower ranked

resources will be added only when the highest ranked resources declines (Winterhalder

1981; Bettinger 1980; Kelly 1995). Ethnographic and experimental research provides general

support for the model (Hawkes, Hill and O’Connell 1982; Hawkes and O’Connell 1985;

Hawkes, O’Connell and Blurton Jones 1991). One disadvantage in applying the model to

archaeological cases is that it requires near perfect paleoenvironmental reconstruction in

order to predict which resources will or will not be used. Nor does it address the proportion

of a resource in the diet as Jochim’s model does. The highly ranked resource may not be

very abundant and encountered infrequently. Its high rank simply means that it will be

taken when encountered.

Kelly (1995) notes the reliance on energy as the sole currency in diet breadth models

may underestimate the value of many resources. Fat and protein content play critical

roles in hunter-gatherer food choices. For example, although the Diet Breadth model

ranks small game such as rabbits lower than deer, because of a lower kcal/kg yield, the

protein/kg of meat in rabbits is nearly equal (Erlandson 1988). Plants, too, may be sought

to provide carbohydrates and fat. In addition, return rates can change due to technology,

seasonal changes in animal behavior and nutritive value, or variations in forager skill

level (Kelly 1995). Seasonal changes in animal behavior and nutritive value are critically

important and will be returned to later.

Patch Choice

The Patch Choice model relies on the assumption that resources are distributed

heterogeneously or in patches and that foragers encounter them randomly (MacArthur

and Pianka 1966; Winterhalder 1981; Kelly 1995). Patchiness can result from the structure

of habitats, prey mobility or resource depression. The patchiness of an environment will

affect the number and types of habitats exploited and mobility patterns (Winterhalder

1981). As with diet breadth, in fine-grained environments foragers will use more patches

and fewer in coarse-grained environments (MacArthur and Pianka 1966; Winterhalder

1981). This is due to increased travel costs between patches. In addition, patches are

ranked according to energy return rates, although in this model search times are included

(Kelly 1995). The highest ranked patch yields the highest energy return rate, but it may

not be the most abundant. Because the model considers search/travel times, widely spaced

patches will lower overall return rates thus resulting in the inclusion of more lower ranked

patches (Bettinger 1991).

Ultimately, the time spent harvesting a patch is determined by the quantity and quality

of resources available (Bettinger 1991). Upon entering a patch the net harvest rate should

be high. As resources are harvested, the return rates decrease. Eventually, search and

pursuit costs outweigh the costs of moving and encountering another patch. Time spent

in a given patch is predicted by a marginal value theorem (Winterhalder 1981; Kelly 1995;

Bettinger 1991). Assuming that foragers will not completely exhaust resources in a patch,

when the return rate of a patch falls below the average for all other potential patches the

marginal value theorem predicts foragers should move to the next patch (Kelly 1995). If

the overall environment is highly productive, foragers will spend less time in a patch and

deplete resources to a lesser degree. Conversely, in lower productive environments,

foragers will spend more time in patches searching for and pursuing lower ranked resources

(Smith 1983).

The model does have some inherent weaknesses. First, the assumption of random

patch encounter is often violated because foragers learn the location of favorable patches

and avoid unfavorable ones. Second, the model assumes travel between patches is

unproductive, which is often not the case. Furthermore, resource density may not fully

predict patch use because while some patches may be rich in certain game, they may be

avoided because vegetation cover increases the search/pursuit costs (Kelly 1995;

Winterhalder 1981). Unfortunately, the data required to test the patch choice model in

archaeology typically are not available. However, Bettinger and Baumhoff (1982) presented

a reasonably successful application that melded the patch choice model with Binford’s

forager/collector model (Bettinger 1991).

Central Place

A third optimal foraging model relevant here is the Central Place Foraging model,

designed to predict the distance foragers will travel to procure resources (Orians and

Pearson 1979; Bettinger 1991; Kelly 1995). The distance is limited by the resource return

rate. Essentially, the model predicts the effective foraging radius based on resource return

rates and the dietary proportion (Kelly 1995; cf. Jochim 1976; Binford 1982). Resources are

ranked according to the ratio of energy to both search time, including round trip travel

time and foraging time, and handling time (Bettinger 1991). The optimal distance a forager

should go is expressed graphically as the intersection of a line representing time drawn

tangential to the expected energy curve. The relationship is similar to the departure time

in the patch choice model, but in this case the intersection is the optimal “energetic ‘cutoff’

point marking the minimal size prey, CI, that ought to be taken given the energetic content

of the patch and the round-trip distance between the patch and the central place” (Bettinger

1991: 93-94). As predicted by Jochim, smaller, easily gathered, reliable resources will be

taken near a camp, while mobile, larger, higher yield resources will be pursued much

farther away. The central place foraging model has the potential to enable archaeologists

to explore the relationships between resource utilization, site function and location as

outlined by Binford (Bettinger et al. 1997). This is important because “subsistence behaviour

at foraging locations cannot be predicted from the quality and abundance of prey alone,

which is assumed in the diet breadth model” (Bettinger et al. 1997: 888). So far, however,

archaeological applications have been limited mainly due to imperfect knowledge of

paleoenvironments and the incomplete archaeological record. These models are currently

being developed and tested using ethnoarchaeological cases (e.g., Barlow and Metcalfe

1996; Bettinger et al. 1997; Bird and Bliege Bird 1997, 2000).


For the most part, the economic tradition of the Cambridge school has been overtaken

by the use of optimal foraging models. The paleoeconomic models developed in the 1970s

and 1980s only considered the caloric yield of a resource. These calculations were based

primarily on meat weight or plant weight and did not take encounter rates or handling

costs into consideration. Site catchment analysis reached its peak in the 1980s when the

approach was largely abandoned due to criticism that landscape use is much more complex

and that use of uniform circles surrounding a given site to estimate an exploitation area

was inappropriate (Limp 1991). Certainly this is true for the early work, however Bailey

and Davidson (1983) took this problem into consideration and modified their territories.

Today, catchment analysis has been “reinvigorated” by GIS application (Limp 1991). GIS

provides a tool for more realistic landscape modeling as opposed to the uniform concentric

circle approach. The main emphasis has shifted from mapping resources near a given site

to estimate potential energy yield to modeling the environment of a large region to predict

where sites should be located given the distribution of various resources.

Without question Binford’s terminology has become deeply ingrained in prehistoric

hunter-gatherer archaeology. In Paleolithic archaeology, this approach to interpretation

is the norm. Optimality models such as Jochim’s and general foraging models are much

more difficult to operationalize, so their explicit use is rare. However, the underlying

assumptions are widely accepted despite existing problems. For instance, Bailey and

Parkington (1988) writing on coastal resource use observed, “(w)hat is at issue here is not

the inappropriateness of optimal foraging theories as such, but of relative cost: benefit

ratios measured under average conditions (or artificial ones), without regard to the

modifying effects of seasonal extremes” (p.6). Jochim’s model was an attempt to get around

the problem by calculating a resource use schedule. In addition, age and sex differences

among foragers need to be considered (Bird and Bliege Bird 1997, 2000; Hawkes et al.

1991, 2001).

Criticisms of the diet breadth and patch choice models focus primarily on the

assumption of energy-rate maximization. Many researchers, especially Jochim, have

argued that reliability or minimizing risk is an important goal in hunter-gatherer decision-

making. Stephens and Charnov (1982) designed the Z-score model with the assumption

of risk minimization. The results were similar to the energy maximization except that

risk-minimizers would leave a patch sooner if the return-rate threshold is greater than the

maximum return-rate and stay longer if the opposite. Winterhalder (1986) applied this

approach to the diet-breadth model with similar results. In this case, diet-breadth should

be restricted when the threshold is greater than the expected return and broader when it

is lesser.

Moderate success in ethnographic cases has led to widespread use of optimal foraging

models in anthropology. Their lack of greater predictive accuracy is likely due to the fact

that they focus solely on energy as currency and consider human behavior in strictly

economic terms. Though optimal foraging models are often criticized by archaeologists

as difficult to operationalize, theorists argue that they are useful as heuristic models for

generating testable hypotheses (Winterhalder 1986).

However, no purely economic model for human behavior can fully explain hunter-

gatherer subsistence and settlement patterns. As Jochim (1998) has written,

…human agency is given paramount importance. This stance allows for the existence
of different behavioral optima and also recognizes that any model of predicted optimal
behavior should be considered as a baseline of expectations, not a description of
real behavior. Optimal models allow us to visualize how people would behave if the
conditions of the models were the only factors affecting their behavior (p.26, italics
in original).

While one could certainly agree with Jochim that human agency should be “given

paramount importance,” energy-based optimal foraging models simply do not allow it.

These models predict behavior as long as the only motivation is energy-maximization.

More recent evolutionary ecology approaches have recognized alternative reasons behind

subsistence decisions (Hawkes et al. 1991, 2001; O’Connell et al. 1998). For instance, Alcock

noted that “(a)lthough some animals do behave in ways that match predictions derived

from this hypothesis, many others compromise energy maximization to deal with such

things as the need for a balanced diet or the need to avoid predators while foraging”

(Alcock 1993: 350). Hawkes (1993) observed that hunters did not pursue large game to

provide food for their families but to acquire status. O’Connell (2000) discussed a similar

pattern among Australian aboriginal hunters. These share the fundamental argument

that natural selection acts upon human behavior that maximizes reproductive fitness, not

necessarily caloric intake (Winterhalder and Smith 2000).

The fact of the matter is that without realistic paleoenvironmental reconstructions,

archaeologists cannot use these models in periods for which there are no modern analogs

because it is impossible to know what resource availability would have been. Without

this knowledge one cannot rank food items and predict what decisions would have been

made. Nor can archaeologists test hypotheses that require observations of subsets of a

given population such as gender or age groups. This is perhaps why most foraging models

are applied to contemporary hunter-gatherers instead of prehistoric ones. Archaeologists

have made a few attempts to build specific models for prehistoric cases. Most of these

economic models consider the potential yield of resources based on modern estimates

extrapolated into the past (e.g. Rowley-Conwy 1984; Straus 1986). More complicated

attempts at patch choice models have been made but they are highly speculative and

usually based on many weak assumptions (e.g., Sept 2001; Zeanah 2000).

Nutritional ecology and human dietary choice

Although specific models for the Paleolithic are rare, the underlying principles of

foraging models are commonly used in archaeological interpretation. The most commonly

held assumption concerns the choice of energy as currency. All paleoeconomic models

assume caloric yield of resources to be the basis for hunter-gatherer subsistence decisions.

A critical problem with the use of energy as the currency is that it ignores broader nutritional

needs. Energy-based foraging models have been criticized for two decades for this reason

(Hill 1987; Keene 1983). Many foraging theorists acknowledge the need to consider other

currencies but most continue to rely on energy because it offers “simplicity” over “reality”,

the latter being too complex (Jochim 1998; Kelly 1995; Sept 2001).

Recently, Kaplan and Hill (1992) ranked resources procured by Aché men and women

separately. Neither foraged according to the predictions of the diet breadth model. Men

ignored plants that offered higher caloric return rates in favor of meat resources. Women

ignored highly ranked animal resources altogether, instead focusing on easily gathered

honey and plants. Kaplan and Hill (1992) write:

Existing optimal foraging models are quite useful for predicting food choice among
resources composed of similar macronutrients but may need to be modified to account
for sensitivity to the nutrient constituents of foods. The assumption that energy is
the sole measure of food value may be inadequate (p.176).

Hockett and Haws (in press) have offered an alternative approach called ‘nutritional

ecology’ to understanding human dietary choice that does not rely solely on energy.

Nutritional ecology is defined as the study of the relationship between essential nutrient

intake and its effects on overall health, including growth and maintenance in individuals

and general demographic trends in human populations. This concept is a slightly altered

definition of that used by Jenike (2001). The nutritional ecology approach is not an

alternative to evolutionary ecology, rather it is based on the same assumption that natural

selection influences human dietary choices. In this case, successful adaptations are those

that ensure a balanced intake of essential nutrients leading to lower infant mortality rates,

greater life expectancy and reproductive success. Nutritional ecology does not measure

foraging efficiency in net energy return of calories from the environment. Instead it focuses

on essential nutrient intake. The model does not assume that foragers will stop to calculate

the nutritional quality of a resource before deciding to pursue it or not.

The nutritional ecology model requires that a number of critical assumptions be

accepted: (1) that hunter-gatherers will make dietary choices that result in balanced diet

whether by accident or intention, (2) that balanced nutrient intake from a wide diversity

of food sources results in better health, (3) that better health leads to lower infant mortality

rates and reproductive success, (4) that lower infant mortality rates and higher reproductive

success are necessary for population growth.

Some argue that hunter-gatherer diets are naturally diverse and they will consume

the necessary nutrients by default. Jochim has argued that foods high in energy are

naturally high in protein and fat and will also provide all of the necessary amino acids,

vitamins and minerals. In many economic models protein content of food items was

considered a sufficient nutritional constraint. However, as Stini (1971) wrote, “(p)roteins

are necessary to supply the necessary balance of essential amino acids and vitamins and

minerals are often not available in adequate quantities in foods that are excellent energy

sources” (Stini 1971: 63). Others, such as Speth, have shown how consideration of

macronutrient (protein, fat and carbohydrates) requirements can alter perceptions about

prehistoric diet choices (Speth 1990, 1991; Speth and Spielmann 1983). From a nutritional

perspective, fats are the best sources of energy, supplying over twice the calories per gram

than protein and carbohydrates (Wing and Brown 1979). The body easily converts fat and

carbohydrates into energy. Protein, on the other hand, can be a potentially dangerous

source of energy. Speth and Spielmann (1983) documented numerous historic cases

illustrating the physiological problems of excess protein consumption. Protein poisoning

will occur when people consume excessive amounts of lean meat in the absence of fat and

carbohydrate because the body will be forced to convert protein into energy to maintain

itself. Fat and carbohydrate are efficient energy sources because both have low specific

dynamic action (SDA). Protein metabolism is inefficient because the body has to break

the protein down and convert the amino acids to glucose to supply the energy it requires.

This leads to greater heat production and energy loss, which must be made up by

consuming even more protein. The SDA for protein is 30%, which means for every 100g

an additional 30g must be consumed. In addition, fat and carbohydrates have what is

called a “protein-sparing action.” When the body uses protein for energy, the first priority

is meeting energy needs, so body proteins are not replenished. In extreme cases the body

must rely on skeletal muscle proteins for energy which leads to dramatic weight loss,

organ failure and, eventually, death. Because protein has a high nitrogen content, this

also leads to increased nitrogen excretion from the kidneys in the form of urea, which also

causes loss of essential minerals and water. The liver can only synthesize a certain amount

of urea each day, and therefore an upper limit of about 50% protein can be safely consumed.

This figure, in fact, may be much lower (Noli and Avery 1988). The sparing action of fat

and carbohydrates reduces the loss of body protein and nitrogen. Studies show

carbohydrates exert a much greater sparing action than fats and thus may be a better

resource during periods of dietary stress ( Speth 1991; Speth and Spielmann 1983). As

Meehan (1977: 493) noted, ‘man does not live on calories alone.’

Besides protein, fat and carbohydrate, humans require many micronutrients such as

amino acids, fatty acids, vitamins and minerals. These are normally ignored in foraging

models but their consideration can also alter perceptions about the value of a resource

and its potential “rank” in prehistoric diets. For several years, paleonutritionists have

attempted to reconstruct the nutrient composition of the “average” Paleolithic diet ( Cordain

et al. 2000; Eaton et al. 1985; Eaton and Konner 1985; Eaton et al.1997). Their work provides

a much better heuristic for understanding prehistoric subsistence. Instead of assuming a

balanced diet was maintained by maximizing energy, their basic questions have been,

“what constitutes a balanced diet?” and “what would a balanced Paleolithic diet consist

of?” Using Lee’s (1968) compilation of hunter-gatherer subsistence, divided into % hunting,

fishing and gathering, Eaton and Konner (1985) and Eaton et al. (1997) produced estimates

for the ratio of plants to animals in Paleolithic diets. They suggest an energy contribution

of 22% fat, 37% protein and 41% carbohydrate based on a plant to animal ratio of 65:35.

Recently, Cordain et al. (2000) revised these figures to a 45:65 plant to animal ratio for

three-fourths of recent hunter-gatherers, based on a reconsideration of the Ethnographic

Atlas data in which hunting and fishing were lumped together. Their conclusion was

that, when possible, hunter-gatherers would maximize meat (balancing protein and fat)

consumption at the expense of plants (carbohydrates). Plant consumption is secondary

but still represents 35-55% of overall diet. Although these ratios are based on recorded

observations of recent hunter-gatherers, biases inherent in the Ethnographic Atlas data

may inflate the economic or dietary importance of certain food groups. Most of the data

come from historic and early ethnographic accounts which are notoriously biased towards

male hunting activities and limited seasonal visits by the observers (Hunn 1981; Milton

2000). In fact, Hunn (1981) shows that the diet among Columbia-Fraser Plateau hunter-

gatherers was around 70% plant foods gathered by women. This alone casts serious doubt

on claims that plants were not important dietary items above 40o latitude. In spite of this,

the ranges in estimated plant:animal ratios and average maconutrient intake show there

may be a fairly high degree of variability in what constitutes an average Paleolithic diet.

Eaton et al. (1998) focused on the importance of long-chain polyunsaturated fatty

acids in human nutrition. Among these are the essential fatty acids, linoleic (LA), linolenic

(LNA), eicosapentaenoic (EPA), docosapentaenoic (DPA), arachidonic (AA),

docosatetrenoic (DTA) and docasahexaenoic (DHA) acid. The latter three are considered

“brain-specific,” that is, they are almost exclusively in the brain and are necessary for

brain growth and development.

Results of nutritional analyses suggest that optimal diets are not simply ones that

maximize energy intake. Nutrient composition should be considered in order to build

optimality models. Balanced diets lead to greater overall health and therefore reproductive

success. Inclusion of food items that may be “suboptimal” in that they lower overall

foraging efficiency may provide nutritional benefits other than calories.

2.1 Explanations for subsistence change

According to the economic optimization models, subsistence change resulting in a

widening of the diet breath or addition of resource patches not previously exploited will

occur when the highest ranked resource or patch declines in abundance or productivity.

Though the Diet Breadth model can be used to determine whether hunter-gatherers are

using “sub-optimal” resources, indices of diversity and evenness are commonly used to

track change through time (Christensen 1980; Cruz-Uribe 1988; Grayson 1984; Kintigh

1988). For the Late Paleolithic and Mesolithic in Europe, Clark and Yi (1983), Zvelebil

(1990), Neeley and Clark (1993) and Bridault (1994, 1997) used diversity measures to argue

for and against increased diet breadth through time. Important factors determining dietary

shifts include climate, technology and demography (Earle 1980). General models for

subsistence change focus on population as the “prime mover” forcing change (Boserup

1965; Christensen 1980; Cohen 1977; Earle 1980).

For the Late Pleistocene/ Early Holocene, Binford (1968) explained the apparent shift

from large game to small game, birds, fish, shellfish and plants as the natural result from

increased population pressure on food resources. As the available land filled with people,

groups were no longer able to move and were thus forced to exploit low ranked resources.

Flannery (1969) formalized this expansion of diet breadth into the ‘Broad Spectrum

Revolution’ to describe the trend in subsistence change in the Near East. He also used

population pressure to explain specialization in large herbivore use followed by

diversification to include “marginal resources” mentioned by Binford. With population

pressure firmly entrenched, Cohen (1977) offered 14 circumstances in which population

pressure could be invoked to explain subsistence change.

Broughton and O’Connell (1999) offer a few methods for detecting resource depression

and subsequent diet broadening in prehistory. First, there is the changing relative

frequencies between large and small game. Second, the level of carcass processing that is

reflected in the damage inflicted on bones. Third, skeletal element representation in relation

to utility could be used “as an index of distant patch use and associated increases in resource

transport costs.” (Broughton and O’Connell 1999: 155).

An alternative explanation for Late Pleistocene/ Early Holocene subsistence change

has been offered by Hayden (1972), who argued that hunter-gatherers maintain controls

to keep population well below carrying capacity so that resource imbalances do not occur.

He later characterized the trend from large to small game as a shift from K-selected to r-

selected resources (Hayden 1981). Instead of population pressure or environmental change,


Table : Prerequisites for population pressure according to Cohen (

) When it is possible to is olate the exploitative cycle of a single group making

its annual round evidence that the range covered is increasing (ie that people
are travelling increasing distances for food) should indicate population pressure

) W hen a group expands into new ecological zone and territories population
pressure may be assumed

) When the inhabitants of a region become more eclectic in their exploitation

of microniches utilizing portions of t he environment such as deserts coastal
areas or forests which have previously been ignored while continuing to
exploit the old niches demographic pressure may again be assumed

) When human populations show a shift toward more and more eclectic food
gathering patterns shown by reduced selectivity in t he foods eaten it may be
argued that they…need to obtain more calories from the same territory in order
to feed denser populations

) W hen a group increases its concentration on waterbased resources relative

to its use of those that are landbased especially …shellfish whose exploitation
is independent of an y new technology this shift may be viewed as resulting
from demographic necessity rather than choice

) When a group shifts from eating large huntable land mammals to eating
smaller mammals birds reptiles and land molluscs demographic stress may
again be assumed

) When a group shifts from the consumption of organisms at high trophic

levels to … lower trophic levels (in p articular when it s hift from meat to plant

) W hen a shift occurs from the utilization of f oods requiring little or no

preparation to foods requiring increased amounts of preparation

) When there is evidence of environmental degradation suggesting human


) When skeletal evidence of malnutrition increases through time

) When the size or quality of individuals exploited from a particular species
shows a steady decline through time (when for example when the size of
molluscs in one or more shell middens decreases)
 ) When an exploited species disappears from the archaeological record it may
be argued that the species was exploited beyond its carrying capacity

 )increasing scarcity of resources should result in the gradual breakdown of the

system of open population flux If this is true then regional specialization of
artifact styles may itself be an indicator of population pressure

) sedentism and the practice of artificial food storage


episodic periods of resource stress led to shifts toward more reliable r-selected resources.

Minimizing risk, not population pressure, is the driving force behind human decision-

making (Hayden 1981; Winterhalder 1986; Yellen 1986). However, Winterhalder (1986)

argued that food sharing is a more effective means of minimizing risk than changes in

diet breadth.

The population pressure model has recently been revised by Stiner et al. (2000). They

argue that instead of a constant population rise culminating in the broad spectrum

revolution, demographic pulses occurred during the Pleistocene as evidenced by shifts in

small game procurement from slow-moving prey such as tortoises and shellfish to fast-

moving lagomorphs and birds. Heavy predation due to population pressure led to over-

exploitation of tortoises and shellfish forcing people to invest more time and energy chasing

rabbits, hares and birds. Over-exploitation was based on size diminution in tortoise humeri

and limpet shells from 4 sites/9 levels in Italy and 2 sites/9 levels in Israel dated between

200-9kya. For Italy, Stiner et al. (2000) show a limpet average size reduction of 1 cm diameter

between the Gravettian (24-28kya) and Early Epigravettian (17-19kya) at Riparo Mochi

dated on lithic typological grounds. In Israel, tortoise humeri size reduction of 0.5 cm

occurs between ~150kya and 100-70kya at Hayonim. Further reduction by 0.2 cm occurs

between the 100-70kya (Middle Paleolithic) and 26-28kya (Aurignacian). Comparison

with the Kebara Cave assemblage indicates size reduction (4.5-4.0 mm) took place between

the late Middle Paleolithic (60-48kya) and Upper Paleolithic (44kya). Stiner et al. (2000)

equate statistical significance tests on limpet and tortoise size reduction with human over-

exploitation due to population pressure.


While the tests reveal significant differences between periods they fail to answer the

question as to how “real” the differences are. Does an average size reduction of 1 cm in

limpet shell diameter or an apparent reduction in tortoise humeri of 3-5 mm reflect “over-

exploitation” or simply “exploitation” (cf. Jones and Richman 1995)? More importantly,

does size diminution reflect “human exploitation?” Claassen (1998) cites numerous

examples of size differences in bivalves, limpets and other gastropods due to natural factors

such as environment, non-human predation, recruitment and juvenile survival, collection

zone/site, and sampling. For instance, juvenile recruitment success and failure has lasting

effects on size frequencies of shellfish, where bad recruitment years can lead to aged or

large-sized dominated populations, whereas one good recruitment year can result in

smaller-sized profiles. Furthermore, many shellfish experience rapid growth over a few

months such that populations exploited during the early Spring may be on average

significantly smaller than those harvested in late Summer (Gordo 1982). Lasiak (1992)

and Meehan (1982) also noted size differences in shells of bivalves and limpets due to

season of collection. Seasonality is therefore a critical factor in assessing measured size

differences in shells through time in archaeological middens.

Recently, Kuhn and Stiner (2001a & b) have pushed the BSR back to the beginning of

the Upper Paleolithic based on the presence of personal ornamentation, namely, the

perforated shells found in the earliest Upper Paleolithic levels of K’sar Akil in Lebanon

and Üçagizli in Turkey. They argue that the need for personal ornamentation was based

on social constraints in the face of demographic pressure. Population ‘packing’ resulted

in the need for group identity and affiliation. Thus, the BSR and population pressure are

tied to the spread of anatomically modern humans into Europe.

The tenets of the nutritional ecology approach lead to a very different definition of

dietary “diversity.” In the diet breadth model, taxonomic richness is used to measure

diversity. Diets based on red deer and rabbit would be considered less diverse than ones

based on red deer, ibex, horse, auroch and rabbit (Grayson and Delpech 1998). The values

in Table 2.2 show that terrestrial mammals have roughly equal proportions of

macronutrients. A similar pattern is seen across fish and shellfish species. Diets that have

one or a few terrestrial mammals provide the same proportions of essential nutrients.

They would not be considered diverse in terms of essential nutrient intake. However,

those that include taxa from the different classes of animals (and plants) provide a truly

diverse range of essential nutrients (Hockett and Haws, in press).

Although terrestrial mammals are excellent sources of energy, they do not provide

all of the essential nutrients required by the human body to maintain itself. Though organ

meats can provide key vitamins not found in muscle tissue, they are often not in appropriate

proportions for human nutritional requirements. For instance, the liver contains vitamins

A, C and E but consuming enough liver to get the necessary vitamin C would lead to

dangerous over-consumption of vitamin A. A much more efficient way of getting the

essential nutrients would be to consume a diverse set of plants and animals. As a

consequence, foraging efficiency would not be measured by energy intake but by dietary

diversity measures across general classes, like mammals, birds, fish, etc. (Hockett and

Haws, in press).

Using the nutritional ecology approach, subsistence change resulting in increased


Table   Comparison of a number of caloric and noncaloric essential nutrients of various

classes of foods All values are based on  g of food
(After Hockett and Haws in press)
c d e f
Essential Terrestrial Terrestrial Shellfish Birds Fish Plants
Nutrient Mammals Mammals
(muscle)a (organs)b
Energy (kcal)  
fat (g)   
protein (g)     

carbs (g)      

C (mg) 

Thiamin (mg)      
Riboflavin (mg)          
Niacin (mg)      
B (mg)       

B (mg) 

A (IU)     
Folate (mg)

D (mcg)  
E (mg)     
Calcium (mg)     

Iron (mg)     

Potassium (mg)   

Based on average values of horse bison red deer rabbit wild boar and
Based on average values of beef liver brains and kidneys
Based on Lamellibranchia
Based on average values of grouse/partridge and duck
Based on average values of Atlantic salmon sea trout and sardine
Based on average values of over  edible plant foods found in the

Mediterranean region 
g of liver alone provides nearly  IU A single daily serving of 

to  IU may be lethal 
Egg yolks contain significant quantities of vitaminD; one egg (the yolk) can
supply  mcg of vitaminD almost as much as g of beef liver

dietary diversity through time is not explained by population pressure on resources forcing

people to adopt energy-poor low-ranked resources. Diets based solely on terrestrial

mammal meat (typically argued for the Middle and Upper Paleolithic in Europe) would

be less efficient leading to less healthy populations, higher infant mortality rates and low

overall reproductive success. Hockett and Haws (in press) have argued that the ability of

anatomically modern human populations to replace Neandertals in temperate Europe

was based on their more efficient, diverse diet. Instead of population pressure based on

some inherent capability to reproduce, anatomically modern humans were able to grow

their populations because their more efficient nutrient intake led to greater reproductive

success, lower infant mortality rates and better overall health (Hockett and Haws, in press).

Chapter 3: A regional perspective on Upper Paleolithic Iberia: a comparison of

Cantabria, Mediterranean Spain and Portugal

3.1 Archaeological applications of economic models in Spain

Archaeologists investigating Late Pleistocene/ Early Holocene subsistence and

settlement patterns in Iberia invariably take an ecological approach. The focus on techno-

environment is made explicit by most researchers (Clark 1991; Straus 1991). Even in the

social and ideological realms, numerous studies have attempted to explain the parietal

and mobiliary art found throughout the peninsula in an ecological framework (Barton et

al. 1994; Clark 1983, 1993, 2000; Conkey 1980; see Mithen 1993 for an alternative view).

Clark (2000) characterizes the portable art as manifesting “assertive” style while parietal

art is evidence of “emblemic” style (sensu Wiessner 1983). Conkey’s (1980) model of

prehistoric aggregation suggests the existence of bounded social groups in the Franco-

Cantabrian region. Despite these attempts, no definitive stylistic boundaries attributable

to ethnic groups within Levantine Spain or Portugal have been identified. Lithic studies

in each sub-region have also failed to discern sub-regional “isochrestic” style zones.

However, there are regional differences in art and material culture between the three sub-

regions that suggest social territories (Arteaga et al. 1998; Straus 1992; Zilhão 1995). The

main basis for delimiting “real” Late Upper Paleolithic culture areas has been the use of

physiographic and ecological features. Within these, archaeologists have reconstructed

exploitation territories or ranges. Different approaches, outlined previously in chapter 2,

have been applied in each area.



Perhaps the best known archaeological sub-region in Iberia is Cantabrian Spain, located

on the northern coast in the provinces of Santander and Asturias (see Figures 1.1 and 3.1).

Bounded by the Cantabrian Cordillera on the south and Bay of Biscay on the north, the

region is divided by river valleys running into the sea and crosscut by rasas, or ridges.

Numerous archaeological investigations have been conducted over the past century due

mostly to the abundance of cave paintings in the region. Over the past thirty years Cantabria

has been the subject of intensive interdisciplinary research designed to reconstruct

subsistence/ settlement systems (Bailey 1983a,b; Bailey and Davidson 1983; Freeman 1973,

1981, 1988; Clark 1983a, 1983b, 1999; Straus 1986, 2000). By far, the works of Geoffrey

Clark and Lawrence Straus have had the greatest impact from a theoretical standpoint.

Their research has shed light on long term trends in subsistence/settlement patterns during

the Upper Paleolithic. Perhaps the most intriguing observation is the gradual diversification

and intensification of the subsistence base from the Solutrean through the Mesolithic (Clark

1987; Clark and Straus 1986; Clark and Yi 1983; Straus 1992). They argued that demographic

expansion and resource stress led to subsistence change as described in the Broad Spectrum

Revolution model. Their argument was based on the faunal record of red deer

specialization and the increase in shellfish use at La Riera cave, which gave support to

Cohen’s (1977) ‘population pressure’ theory (Clark and Straus 1986). In their test of the

Cohen model, the sole criterion met was size reduction of a particular species, in this case,

limpets. Subsequent research has continued the use of the BSR model (Straus 1991, 1992,

1995a, 1995b, 1996; González Morales 1995; González Sainz 1995; Aura et al. 1998; but see

González Morales 1991for a different view). Here, as in other areas, exploitation of a

diverse range of plants and animals coincides with the spread of forests back into the


Cantabria was a fertile testing ground for site catchment analyses in the 1970s and

1980s (Bailey 1983; Bailey and Davidson 1983; Clark 1983; Clark and Lerner 1980). Clark’s

(1983; also Clark and Lerner 1980) work focused mainly on Mesolithic subsistence and

settlement patterns. Bailey and Davidson (1983) made explicit use of site catchment and

site territorial analyses for Cantabria between 20,000 and 15,000 bp, corresponding to the

Solutrean and Lower Magdalenian. Bailey (1983) grouped the Cantabrian sites into clusters

that share the same site exploitation territory based on 2-hour distance radii creating tightly

packed site yet mutually exclusive territories along the Cantabrian coast. The three main

site clusters, El Castillo, La Riera and Lloseta, are located in good positions to exploit

interior/upland ibex, coastal plain red deer, and coastal shellfish resources. Bailey argues

these three represent major site systems capable of providing resources year-round.

However, one of the main difficulties in reconstructing subsistence/settlement systems

for this period is that the Last Glacial Maximum (LGM) coastline would have extended

an additional 8-10 km, meaning a significant portion of the archaeological record is likely

submerged under the sea (Clark 1983; Perlman 1980). An alternative view proposed by

Craighead (1995) suggests an even shorter distance.

Straus and Clark (1986) used site catchment analysis and the forager/collector model

to reconstruct subsistence and settlement patterns. Straus’ (1986) model was illustrated

El Juyo
La Riera
N Altamira Rascaño

Cingle Vermell
Atlantic Ocean

Cova Fosca
Buraca Grande
Lapa do Picareiro Caldeirão Cova dels Blaus
Bocas Cueva de la Cocina
Lapa do Suão
Volcán de Faro
Les Mallaetes
Parpalló Santa Maira
Figueira Brava Tossal de la Roca Cova de les Cendres

Vale Boi


Vanguard Cave Mediterranean Sea

Gorham's Cave

Figure : Map of Iberia showing the main sites mentioned in the text

by drawing foraging radii around residential sites with arrows pointing to the hilly and

montane regions where ibex and other resources were exploited. Straus argues for a

logistical exploitation strategy due to “optimally situated base camps” and high resource

density (Straus 1986: 331). Site location seems best determined by ‘blocking’ locations in

river valleys where red deer can be effectively hunted suggesting a specialized adaptation

(Bailey 1983; Straus 1986).

The best attempt to quantify the available or potential resources for Late Upper

Paleolithic Cantabria is Straus’ (1986) model, which sought to determine the number of

microbands supportable by red deer in the Solutrean and Magdalenian. His approach

was similar to Jochim’s in that it was based on the estimated red deer carrying capacity by

analogy with the present-day Scottish Highlands. Following Vita Finzi and Higgs, Straus

drew upon Darling’s “Herd of Red Deer” study from the Scottish Highlands in the 1930s.

In many ways , Darling’s study is more applicable to Upper Paleolithic northern Spain

than the Levant due to environmental similarities between present-day Atlantic heathlands

of Scotland and Late Glacial Maximum Cantabria (Straus, 1986). Darling gave an average

of one red deer per 40 acres for the Highlands, with a range of 1 per 30 acres to 1 per 100

acres. In total, he counted 1315 deer in the 52,000 acre area. Population balance was

maintained by a 20% annual cull combined with a 50% mortality rate in the first year. The

average weight of the adults ranged from 125-200 kg. After calculating the total land area

during the LGM (1600 sq. km), Straus estimated red deer populations for eastern Asturias

between 8-10,000 (80-100 herds of 100 animals). Assuming an average of 100 kg usable

meat per animal a 10% cull rate, red deer could have provided 8000 kg or 320kg/person in

a 25 person band. With a 1400 kcal/kg average venison yield and a cold-climate dietary

requirement of 3,600 kcal/day/person for the LGM (a person would need to eat 2.5kg per

day!), Straus proposed 8-10 bands of 25-30 people for eastern Asturias during the LGM.

Although overall population density may have been low, coastal densities were likely

much higher (Straus 1986). While this model assumes exclusive consumption of red deer

it serves to illustrate the potential resource availability. Straus rightly estimated the

extraordinary amount of lean meat that would have been required to avoid protein

poisoning in the absence of fat and carbohydrate.

However, several problems arise with these estimates. Straus uses plane surface area,

not taking into consideration topographic variation. As alluded to earlier, the topography

of Cantabria is cut by river valleys, hills, ridges, and mountains making use of planar

surface area suspect in estimating animal populations. Another problem is that in the

Scottish Highlands the herds of red deer are managed for sport hunting. All predators

except humans have been eliminated, as have competing animals. While it is true that

archaeofaunas are dominated by red deer and ibex, it is assumed that this represents the

natural background availability. This may be true, however it must be realized that site

location seems best determined by points on the landscape where red deer can be most

effectively hunted (Bailey 1983; Straus 1986). This suggests a specialized adaptation. What

we ‘see’ archaeologically may be clouded by human decision-making, a problem in site

catchment analyses (Davidson 1983). An additional problem is the use of meat weight

estimates for red deer in order to calculate the potential human population. No one,

including Straus, would argue that hunter-gatherers were consuming red deer meat

exclusively, but the model becomes inherently flawed when the assumption is made, in

spite of its simplicity. Even used as a heuristic this approach will not accurately estimate

population, although there appears to be a desire among archaeologists to attempt it.

Mediterranean Spain

The coastal strip of Spain from Catalunya down to Valencia and across Murcia to

Andalucía has been a focus of paleolithic archaeologists for decades (see Figures 1.1 and

3.1). This region is often referred to as Levante or the Spanish Mediterranean Region

(SMR). It is quite different than Cantabria in geography and environment. In constrast to

the northern coast, the Mediterranean coast is characterized by a broad coastal plain which

extended 20-30 km further during the Late Pleistocene. This plain is bounded by mountains

with elevations up to 3000 m, making the region geographically circumscribed. The

vegetation is strictly Mediterranean and is discussed further in chapter 4.

Economic life in the Upper Paleolithic was described by Pericot (1968). He recognized

that Paleolithic hunter-gatherers depended heavily on meat from large game but also

consumed a wide variety of small game, birds, fish, shellfish and plants. He argued that

shellfish were as important to people who occupied Parpalló as abalone was to prehistoric

coastal peoples in California. For plants, the most likely ones included were hazelnuts,

acorns, walnut and chestnuts. Additionally, wild fruits, berries, roots, tubers, grass seeds,

fungi and even seaweed were likely important dietary items (Pericot 1968).

The earliest attempts to reconstruct subsistence and settlement patterns were done

by Davidson (1976, 1983) and Bailey and Davidson (1983). Both studies undertook site

catchment and site territorial analyses for the Late Pleistocene in this region. Both used

faunal and artifactual remains to estimate the site exploitation territories of Upper

Paleolithic sites in Valencia. In general, these sites, which contain deposits dated to the

Gravettian and Solutrean, are dominated by rabbit, red deer and ibex.

Davidson (1976) used the 2-hour and 1/2-hour walk radius of Vita Finzi and Higgs

(1970) to calculate site exploitation territories for the Upper Paleolithic sites Parpalló, Les

Mallaetes in the Mondúver massif and Volcán de Faro on the coast of València. Examination

of Parpalló and Les Mallaetes reveals some important differences between the two.

Although they share much of the same exploitation territory (79% overlap) they may be

different seasonal base camp locations. Parpalló, located at 450m asl, has a south facing

entrance whereas Les Mallaetes, 600m asl, has a west facing entrance, exposed to prevailing

winter winds. Parpalló has better access to low elevation flat pastureland in an area beyond

the overlap with Les Mallaetes. On the other hand, Les Mallaetes has better access to

higher elevation flat areas. Although the two certainly have access to ibex and red deer

habitats, Parpalló would have been better suited to hunt red deer in the winter and Les

Mallaetes to hunt ibex in the summer (Bailey and Davidson 1983). Indeed, Parpalló has

numerous hearths and an overabundance of antler while Les Mallaetes has little antler,

male ibex (presumably on their own during the summer), neonates, and few hearths

(Davidson 1983).

Davidson followed this line of thought further by experimenting with 1/2 hr mutually

exclusive site exploitation territories. Within this area there are seasonal advantages.

Parpalló, however, is not situated so much for access to grazing areas as it is advantageously

located in a valley corridor to hunt red deer on their migration to and from summer

pastures. Given the antler evidence for site seasonality, it may have been occupied when

red deer were leaving the upland pastures. Turning to the coastal lowlands, Davidson

suggests that the Maravelles and Porcs sites were nearer to lower elevation pastures that

may have been exploited in winter. Thus, there may have been a pattern of seasonal

transhumance between the coast and the upland interior. Marine shells have been found

at Parpalló and Les Mallaetes which were about 30 km from the LGM coast. It would

seem unlikely, based on this, that the economic catchment and the site exploitation territory

were the same during this period. However, similar red deer to ibex ratios for Les Mallaetes

and Parpalló led Davidson to favor the 2-hour radius for site exploitation territories, which

allows for access to both resources. He put forth climate change as a possible explanation

for why these two would have similar ratios. It seems unlikely that this could explain the

similarities since they were both occupied during the Last Glacial Maximum. The idea of

a large (2 km) exploitation territory makes more sense.

At Volcán, the 2-hour site territory contains both red deer and ibex habitat. Within

the 1/2-hour territory, ibex habitat dominates the area, but the abundance of red deer

contradicts notions that it was ideally suited for ibex exploitation (Davidson 1983). Summer

occupation might have reduced the likelihood that ibex were present near the site and

increased the availability of red deer. If this is truly the case, then a model of transhumance

may not be accurate. There may have been two groups living side by side, one in the

interior and the other on the coast with some interaction apparent between the two. Though

Davidson (1983) suggested this possibility his acceptance of the larger exploitation

territories for the two interior sites would make it unlikely. A third possibility is that the

resolution is too coarse-grained and that Parpalló and Les Mallaetes were occupied in

alternating years or generations. Such a case would not show up with the limitations in

dating methods for the Upper Paleolithic.

As Straus did for Cantabria, Davidson used meat weight estimates to model the

subsistence economy for Mediterranean Spain. The main difference between the faunal

record in Mediterranean versus Cantabrian Spain is the extreme abundance of rabbit

remains in the former. Rabbits have always been problematic in Iberian cave and

rockshelter assemblages due to their burrowing habits and deposition by predators such

as lynx, fox, and eagle owl. Recent taphonomic work by Hockett, Pérez Ripoll and Serra

in Iberia has shown differences in characteristics of assemblages deposited by humans

and predators. These will be discussed later in chapter 5.

Davidson (1976) assumed for the sake of modeling that people deposited most of the

rabbit bones in the caves and rockshelters of Mediterranean Spain. He then addressed the

problem of the importance of rabbit in the diet during the Upper Paleolithic. Using dressed

meat weight of rabbit, red deer and ibex he concluded 100-150 rabbits would be necessary

to equal the amount of meat from a single adult male red deer. Based on MNI estimates

from Les Mallaetes he argued rabbit comprised less than 3% of the diet. Furthermore,

using estimates of rabbit hunting rates from modern ecological studies he calculated a

hypothetical rabbit-based economy. Davidson used 1957 FAO figures of 2,750 cal/day as

the mean human requirement and an energy estimate of 120 kcal/100g for rabbit meat.

Using an estimate of 7.41 rabbits per hectare, based on modern Australian and British

rabbit densities, and a 30-40% cull rate, a 7,300 ha area would be necessary to feed 20

people per year. This area was within the 2-hour walk radius of both Parpalló and Les

Mallaetes. Unlike Straus, Davidson anticipated optimal foraging concerns over hunting

success rates. Again using modern rabbit control methods as a baseline, he estimated a

0.5 rabbits per hour procurement rate for the Paleolithic, which would mean the same 20

people would have to spend five hours per day hunting to subsist exclusively on rabbit

(Davidson 1976). Adding the 2 hours’ walk each direction this would actually require 9

hours per day! Given these estimates, Davidson dismissed the possibility that rabbits

were an important component of Upper Paleolithic subsistence in Spain. Obviously, rabbits

were not the sole source of food as this model assumes.

Davidson, Bailey and Straus all calculated the necessary amount of meat from a single

species needed to fulfill daily caloric requirements. These single species-based hypothetical

economies were used to estimate human population and evaluate dietary importance of

resources. Each used FAO daily caloric requirement estimations. Straus modified these

to account for the rigorous climatic conditions of the LGM. Bailey (1978) and Davidson

(1976) used meat weight estimates to “test” small animal-based economies. These two

conclude that neither rabbits nor shellfish could have been important regular components

of prehistoric diets because to the extraordinary high numbers of each animal required to

feed a person. For rabbits, hunters would have to capture 100-150 to equal the amount of

meat from a single red deer. For shellfish, 1,000 cockles would be necessary to meet a

single person’s daily caloric requirement, 150,000 to equal the meat from a single red deer.

However, the assumption is that these single animals would be the sole food source.

Furthermore, if either resource cannot reasonably sustain people for a day or year because

the procurement costs would be too high or the sheer volume of meat necessary would be

unattainable then they could not rank very high. In that case, despite the apparent

abundance of their remains in the archaeological record, they should be ignored or

considered as a biased numerical over-representation, a mere side note.

These models are intended to serve as heuristic devices to understand past subsistence

behavior. Single resource dominance in human diet is rare and usually occurs in extreme

environments, if ever. Even in the French Upper Paleolithic reindeer did not comprise

100% of the diet. Its abundance may be a seasonal reflection, whether the “herd-follower”

model is preferred or not. Furthermore, even though Davidson and Bailey show low

meat weight percentages for rabbit and shellfish, ranking them low in the overall diet, the

undeniable fact is that their remains do not just dominate assemblages, they swamp them

in many cases. People went to some effort to procure them on a large scale. Why? Foraging

model expectations and the BSR model would imply they were under stress from

population/resource imbalance. However, as ethnographic studies have shown, resource

rank does not always correspond to dietary proportion (Hawkes et al. 1991, 1997, 2001;

Kaplan and Hill 1992). A broader nutritional perspective and/or one that considers women

and children would provide a different answer. Red deer and ibex may have been highly

valued resources for reasons other than provisioning hunters and their families. Rabbit,

plants and shellfish being more easily gathered may have constituted the ‘daily bread’.

Since the work by Davidson, a new generation of Spanish archaeologists have


continued the lines of research he set forth. The biggest revolution in Spanish archaeology

has been the widespread focus on taphonomy. This field has especially grown with the

excavations at the Sierra de Atapuerca where the largest collection of archaic human

remains in Europe were unearthed and the Orce basin where the oldest hominids in Europe

have been found. In addition the late dates for Neandertals in Iberia have necessitated

detailed taphonomic studies of the caves and rockshelters yielding Neandertal remains

and evidence of human behavior across the Middle-Upper Paleolithic transition. Current

research suggests Neandertal and modern human populations lived in close proximity

separated by the Ebro River in northern Spain for 10,000 years before modern forms

replaced the Neandertals. The quality of these studies has led to an acceptance on the part

of most European prehistorians that the Iberian Peninsula is a critical area to study the

Middle-Upper Paleolithic transition and the dynamic interaction between two hominid


Emphasis on human behavior and taphonomy instead of typology and culture history

has carried over into all aspects of Iberian prehistory. However, the theoretical framework

for explaining past human behavior has not expanded beyond that provided by Vita-

Finzi, Higgs and Binford. General explanations of Upper Paleolithic subsistence and

settlement patterns since Davidson’s work are based on the fundamentals of site catchment

analysis, using meat weight as a measure of dietary importance and the Broad Spectrum

Revolution model. The Davidson (1989) model of seasonal coastal-inland movement for

Upper Paleolithic settlement patterns has been strengthened with data from additional

recently excavated sites. This is discussed further in chapter 5.


With regard to meat weight and estimating relative proportion of rabbit, shellfish

and medium ungulates, Villaverde and Martínez Valle (1995) reiterate Davidson’s (1976,

1989) point that to get the same amount of kilocalories from a single red deer or ibex, one

would have to procure 100-150 rabbits. Morales et al. (1998) provide a similar estimate as

Bailey (1978) did for shellfish. Villaverde and Martínez Valle (1995) also characterize

rabbit as a nutritionally poor resource for human consumption because of its low fat content.

However, as Table 2.2 shows, rabbit contains more grams of protein and fat than red deer

and in more balanced proportion. The main difference is package size.

The methodological developments in Americanist zooarchaeology beginning with

Binford (1981) have been almost completely ignored by most Spanish researchers. The

sole exception is the work of Maria Fernanda Blasco Sancho (1995, 1997). Almost all faunal

tables in site reports from the region present data in NISP. Occasionally, MNI are reported.

Where skeletal element representation is considered, element frequencies are calculated

by NISP, not MNE. The problems with this approach are discussed in further detail in

chapter 5.

Following Davidson (1976, 1989) several authors have argued for a trend towards

resource intensification and diversification at the Pleistocene-Holocene transition (Aura

Tortosa and Pérez Ripoll 1992, 1995; : Aura et al. 1998, 2001; Aura 2001a&b; Pérez Ripoll

and Martínez Valle 2001; Villaverde and Martínez Valle 1992, 1995; Villaverde et al. 1998;

Villaverde 2001). Aura and Pérez Ripoll (1992) initially argued that the high degree of

rabbit exploitation in the Magdalenian was a sign of a broadened dietary spectrum. As

Villaverde and Martínez Valle (1995) pointed out this level of exploitation is apparent as

early as the Aurignacian in sites like Cova Beneito. Prior to Stiner et al. (2000), Villaverde

and Martínez Valle (1995) considered differences between small game types. The changes

in diet breadth at the end of the Pleistocene were not simply an inclusion of small game in

general. Rather, faster moving, harder to catch small animals such as hares and birds

were argued to signal greater dietary diversification as predicted by the Broad Spectrum

Revolution and diet breadth models (Villaverde and Martínez Valle 1995). Villaverde et

al. (1997) proposed a model for small game based on reduced mobility. Because rabbits

are abundant and territorial, hunter-gatherers could have economically exploited them

by reducing residential mobility. Thus, people may have become tethered to small game

and possibly plants at the beginning of the Upper Paleolithic.

Summary faunal tables for Cova de les Cendres based on NISP do show the rise in

frequency of hares and birds in the Late Upper Paleolithic (Villaverde and Martínez Valle

1995). However, the numbers are extremely low: NISP of 10 hare bones in the Upper

Solutrean to 45 in the Middle Magdalenian to 19 bones in the Upper Magdalenian. At

Cova dels Blaus, hare NISP increase from 122 (13%) in the early levels V & IV (Upper

Magdalenian/ Microlaminar Epipaleolithic) to 287 (16.6%) in level III (Microlaminar

Epipaleolithic) to 92 (16%) in levels II & I (Villaverde and Martínez Valle 1995). The rise in

the proportion of hares is very slight and does not appear to represent a significant dietary

shift across the Pleistocene-Holocene transition. Aura Tortosa and Pérez Ripoll (1992)

show an actual cumulative decrease in birds from the Upper Magdalenian to the Geometric

Epipaleolithic. Total NISP for birds in Mediterranean Spain is 537 (2% of the total) in the

Upper Magdalenian, 285 (2%) in the Microlaminar Epipaleolithic and 14 (0.7%) in the

Geometric Epipaleolithic. Again, this casts doubt on the use of hares and birds as indicators

of a diachronic trend towards diversification for this period. A longer perspective including

the Gravettian and Solutrean suggests some validity due to the absence of hares from the

sites in Mediterranean Spain. A variety of birds (at least 18 species) and fish are present in

the Solutrean levels of l’Arbreda (Éstevez 1987; García Petit 1995, 1997; Muñoz and

Casadevall 1997). The highest frequency and greatest diversity of fish were found in the

Gravettian and Solutrean levels. Fish numbers decreased significantly in the level dated

to the coldest period of the LGM (Muñoz and Casadevall 1997).

Morales et al. (1998) have succinctly characterized the consensus view of Late Upper

Paleolithic subsistence in Iberia as the ‘Tardiglacial paradigm,’ namely that resource

intensification and diversification began sometime during the Solutrean. In their study of

marine resources at Cueva de Nerja, the primary features include:

(1) a diversification of subsistence strategies through time

(2) an increasing reliance on aquatic resources
(3) evidence of the progressive approach of the coastline which, after a
threshold being reached, would allow the cave dwellers to exploit resources
which were previously too far away for systematic cropping
(4) an indication of changes in settlement strategies which, among other
things, meant that resources which were previously consumed in open-air
camps at the seashore started having wider patterns of distributions
throughout the areas under study (Morales et al. 1998: 41).

Unfortunately, there is no definitive Solutrean occupation of Nerja and the Aurignacian

and undifferentiated Early Upper Paleolithic levels contained very few faunal specimens

(Jordá 1986; Morales et al. 1998). Therefore, conclusions about diachronic trends are

untestable at Nerja.

Despite the inter-site variability and the differences in the nature and timing of the

inclusion of small animal resources in the diet, a consensus has been reached among Spanish

archaeologists that resource intensification and diversification characterize subsistence

changes in the Late Upper Paleolithic and Epipaleolithic archaeological record of

Mediterranean Spain. In contrast with Straus and Clark for Cantabria, the suggested

mechanism for this development is not demographic pressure (Aura and Pérez Ripoll

1992; Aura et al. 2002). Higher regional population is hinted at by the expanded use of

upper elevations and increased number of sites, but it is not explicitly stated as a causal

mechanism. Instead, reduced mobility and an integration of rabbit, and presumably, other

resources into hunter-gatherer economic systems after the appearance of modern humans

is the reason given.

The work in Cantabria and Meditearanean Spain has had an important impact on

research in central Portugal. Many of the assumptions from the meat weight studies and

the Broad Spectrum Revolution model have been incorporated into models developed by

archaeologists working in this region. These studies are discussed in the following section.

3.2 Central Portugal

Portuguese Estremadura is a coastal region of approximately 8,700 km2, located

between 40° and 38°30' north latitude, encompassing the political provinces of Estremadura,

Ribatejo, and part of Beira Litoral (Figure 1.2). It is bounded to the north by the Mondego

River and to the south by the Tejo River, though some argue it extends to the Sado River.

On the eastern boundary lies the Serra da Estrela, a schist and granite mountain rising up

to 2000m. The region is characterized by a limestone massif, rising up to 650m, with

maquis or garriques vegetation and karstic caves (Zilhão 1990). Natural vegetation is a mix

of Atlantic and Mediterranean species, with the latter dominant. Estremadura is

interspersed with montane islands surrounded by plains and valleys, under continuous

cultivation of olive, wine grape, and fruit trees. Eucalyptus plantations are rapidly replacing

pine forests of the region. Deep plowing for eucalyptus has greatly benefited archaeological

survey for open-air paleolithic sites.

Recent models

Table 3.1 shows the chrono-stratigraphic framework for the Portuguese Magdalenian.

In general, this period is characterized by microblade tool production based on a variety

of core types. The chronological divisions of the Magdalenian are based on differing

frequencies of these types through time.

For the Upper Paleolithic of Portugal, Zilhão (1990, 1995), Bicho (1994, 1996), Thacker

(1996) and Shokler (1995) have proposed models based on lithic technology and raw

material economy, distribution of sites, and limited faunal data. Each model suffers from

a lack of crucial data on subsistence and seasonality. Until the excavation of Lapa do

Picareiro, reconstructions of Upper Paleolithic subsistence relied on paleontological

collections, taxonomic lists from sites excavated in the late 19th and early 20th centuries,

and a few unpublished or partially published sites excavated in the last few decades.
Table : Late Upper Paleolithic and Epipaleolithic chronostratigraphy after Zilhão (

Early Magdalenian Cabeço do Porto 

 bp Various types of cores including prismatic
(CPM facies) Marinho (CPM) bladelet cores burins on flake or cortical
Caldeirão blanks but few on thick scrapers; backed
microliths are  backed bladelets

Early Magdalenian Cerrado Novo  bp Prismatic and thick endscraper cores near
(Cerrado Novo facies) Vascas absence of cores on burins; microliths 

Dufour bladelets Areeiro bladelets and
marginally backed bladelets
hiatus   bp no sites dated to the Middle Magdalenian
Upper Magdalenian Buraca Grande   bp similar to CPM facies; microliths 
CPM Caldeirão bladelets and backed points most on burin
Picareiro Vale da spalls
Mata Vascas Suão
Final Magdalenian Pinhal da Carneira  bp cores mainly on burins; microliths  
(Rossio do Cabo facies) Rossio do Cabo Dufour bladelets Areeiro bladelets and
marginally backed bladelets
Final Magdalenian Bairrada Bocas  bp significant blade production; bladelet cores
(Carneira facies) Carneira Olival da mainly on burins; microliths  geometrics
Carneira Picareiro (trapezes and segments)
Boreal Mesolithic 
 bp cores mainly on thick scrapers; microliths

 Dufour bladelets Areeiro bladelets and
marginally backed bladelets

Despite these limitations each model uses a Binfordian framework for describing the

nature of settlement patterns. Though few sites in Estremadura fulfill the expectations of

a residential camp based on size, studies of lithic assemblages by Zilhão and Thacker led

to the classification of a number of sites as ‘residential base camps.’ For the Magdalenian,

Zilhão (1997) considers the open-air sites Bairrada, Cabeço do Proto Marinho (CPM),

Cerrado Novo and Olival da Carneira to be ‘base camps’ (Figure 3.2). This was based on

their location in strategic places on the landscape and the occurrence of a high diversity of

tool types in all stages of manufacturing. Unfortunately, none of these contained preserved

faunal remains. In contrast, none of the fauna-bearing sites could be classified as residential

sites. All are in small caves or rockshelters that were likely used as hunting camps or

butchery/processing sites. However, Zilhão (1995) has suggested Caldeirão functioned

as a base camp throughout the Upper Paleolithic. The lack of long-term residences does

not mean that hunter-gatherers during the Late Pleistocene employed an exclusively

“foraging” strategy. Considering lithic raw material procurement and tool manufacturing

during the Magdalenian, Thacker (1996, 2000) argued in favor of a foraging pattern.

Whereas flint cores were prepared at lithic quarry sites during the Gravettian, whole cobbles

were brought to sites 1-2 km away during the Magdalenian. Furthermore, Thacker (2000)

argues that the decreases in local quartz and quartzite use suggests greater residential

mobility during the Gravettian than the Magdalenian since base camps would contain a

wider range of raw material if they were occupied for long periods. The fact that

Magdalenian assemblages are dominated by flint as opposed to Gravettian ones may

suggest a collector pattern for the latter and a forager one for the former. On the other

Buraca Grande
Atlantic Ocean

Costa do Pereiro
Lapa dos Coelhos

Bocas O de Carneira
Lapa do Suão

Rossio do Cabo
Cerrado Novo
Vale da Mata





Figure  : Magdalenian sites in Portuguese Estremadura


hand, Bicho argues for a collector settlement pattern during the Final Upper Paleolithic

and Epipaleolithic simply because specialized task sites appear more frequently. The

presence of specialized task sites may not necessarily indicate greater logistical mobility if

these fall within the foraging radius of a residential site.

The analysis of CPM V and Carneira, two open-air sites in the Rio Maior valley, showed

variability in raw material use during the Late Magdalenian (Bicho 1994). Carneira, located

less than 2 km from flint sources contained 95% flint while CPM V, located 2.5 km from a

source, had about 75% with local quartz and quartzite making up the rest. This suggests

that distance to flint sources influenced decision-making. According to Zilhão (1997),

Carneira was probably an intermittent camp while CPM was likely a base camp. The data

on lithic raw material from each agrees with Thacker’s assertion that base camps should

have greater raw material diversity.

Zilhão (1995, 2000) has drawn on the Selk’ nam or Ona of Tierra del Fuego as a source

of analogy for central Portugal during the Late Pleistocene. He argues that this is

appropriate because the Ona occupied an environment analogous to Portuguese

Estremadura during the Last Glacial Maximum and a territory similar in size. They hunted

a medium ungulate and small rodent roughly equal in size to red deer and the European

rabbit, ubiquitous in the Upper Paleolithic of Portugal. The model uses carrying capacity

estimates derived by Straus (1986) for Cantabria and Bailey et al. (1983) for Epirus. Using

their estimate of 0.05 people/km2 and a minimum band size of 25, there may have been

20-24 microbands in Estremadura with territories of 400-500 km2.

Zilhão (1995) also noted a diachronic shift in settlement patterns between the

Gravettian and Magdalenian in Estremadura. Both industries are characterized by

retouched microliths. The site types discernible in the Gravettian include specialized

workshop sites, specialized logistic sites and generalized residential sites. However,

Magdalenian assemblages exhibit a greater homogeneity leading to the conclusion that a

forager settlement pattern was in place. This was also argued by Thacker (2000).

After 16,000 bp, when climate appears to have rapidly ameliorated, Magdalenian

hunter-gatherers began specializing on red deer and rabbit. According to Lee (1969), who

relied on Murdock’s Ethnographic Atlas, the Ona derive 80% of the diet from animals: 60%

terrestrial game, 20% fishing. Therefore, one could argue that while plants were utilized

in both cases, their overall dietary contribution was minimal and the appropriate focus

for subsistence study is on animal resources.

Zilhão (1997) has divided Estremadura into three hypothetical territories based on

river drainages, site clusters and symbolic differences. These are based on three (and

possibly four) site clusters in Estremadura: the Cambelas area immediately north of Lisbon

near the city of Torres Vedra including the Rio Maior area at the foot of the Serra de

Candeeiros, the Massif area encompassing the Serras de Aire and Candeeiros, and Mondego

river area on the northern edge of Estremadura and bounded by the Serra do Sico on the

east. Each area encompasses approximately 4,000 km2 which is roughly equivalent to the

Tierra del Fuego case. Each exhibit the same subsistence regime and technology during

the Upper Paleolithic. Perforated shell ornaments from Littorina obtusata are prevalent

and there is limited evidence for red ochre. The primary reason for the delineation, other

than the clustering which is almost certainly artificial, is the presence of perforated red

deer phalanges at Caldeirão in the Massif area an their absence to the south and north.

However, the bilaterally perforated red deer phalanges could be interpreted as examples

of carnivore tooth punctures. Thus, there may be no real basis for delineating cultural or

paleoethnographic boundaries in Estremadura using the presence or absence or portable


Bicho model


Another model of subsistence and settlement during the Final Upper Paleolithic and

Epipaleolithic of Portugal comes from syntheses by Bicho (1993, 1994, 1996) and Bicho

and Haws (1996). Paleoenvironmental changes during the Tardiglacial were relatively

minor but still induced fluctuations in the availability of ibex, chamois and red deer.

Between 15,000 and 10,500 years ago, a general warming trend led to the replacement of

ibex and chamois by red deer, wild boar, aurochs, and rabbit in the faunal assemblages of

Caldeirão (200 m asl) and Lapa do Picareiro (500 m asl). The recovery of chamois at

Picareiro suggests elevational changes in the biogeographical distribution of this species

as a result of climate change, which seems to be confirmed by its absence at Caldeirão. By

10,500 bp, diversification is apparent in the addition of marine and estuarine shellfish

found in coastal middens as well as caves 35- 50 km from the coast. Shell middens are

known from Gruta do Casal Papagaio, Pena da Mira, Bocas, and Cabeço de Curral Velho

(Araújo 1995; Arnaud and Bento 1988; Arnaud 1993; Bicho 1993b, 1994b). Perforated

ornamental shells and edible species of bivalves have been recovered from contemporary

Buraca Grande
Atlantic Ocean

Casal Papagaio
Pena de Mira Pena d'Agua

Areeiro III
Lapa do Suão

Ponte da Vigia

Cabeço do Curral Velho

São Julião

Penha Verde Lisboa

Quinta da Bicuda
Ponte de Cabadelo


Figure  : Epipaleolithic/Early Mesolithic sites in Portuguese Estremadura


levels at Lapa do Picareiro (Bicho and Haws 1996). Fish has been identified at Lapa do

Suão, Caldeirão, and Picareiro (Bicho 1994b; Roche 1979, 1982; Zilhão 1992a, 1992b, 1993).

The coastal shell middens of Toledo, Magoito, São Julião, located in sand dunes, have

been dated to the Pleistocene-Holocene boundary (Bicho 1994b; Daveau et al. 1982). A

surface scatter of Epipaleolithic lithics and mollusc shells has been documented at Quinta

da Bicuda, but the association is in doubt (Carvalho, 1996). There are also shell middens

known from Alentejo (Pedra do Patacho) and Algarve (Castelejo) that date to this period

(Bicho 1994b; Bicho and Haws 1996; Soares and Da Silva 1993; Vierra 1995).

In summary, dietary data suggested similarities between central Portugal and northern

Spain. The observed pattern in both regions is characterized by gradual subsistence

intensification through specialization and diversification at the end of the Pleistocene

(Hayden 1981). Specialization was based on the presence of hunting sites with faunal

assemblages dominated by red deer and aurochs (Bocas), and rabbit (Picareiro). However,

it is not likely to have been as important as it was in northern Spain (Bicho and Haws 1996;

Straus 1992, 1995). During the Preboreal and Boreal (10,000-8,000 bp), an increase in dietary

diversity is seen by the inclusion of birds and aquatic resources such as fish, cockle, oysters,

clams, mussels, limpets, and top shell. Besides animal resources, plant foods such as

fruits, nuts, seeds, and berries were likely used. The appearance of grinding stones in the

Gravettian and Magdalenian suggested that dietary diversity was an important component

of hunter-gatherer adaptations throughout the Upper Paleolithic (Bicho and Haws 1996;

Hockett and Haws 2002b).


According to the model, there are two discernible phases of settlement patterns in

Portuguese Estremadura. The first phase corresponds to the Early, Middle and Late

Magdalenian, while the second phase corresponds to the Epipaleolithic, starting around

10,500 bp (Figures 3.2 & 3.3) (Bicho and Haws 1996).

During the first phase, there were three major areas of occupation, two inland near

Rio Maior and Torres Novas, and one coastal near Torres Vedras. These are equivalent to

those outlined by Zilhão (1997). Sites appear to be preferentially located on the flat plains

found in the river valleys. These areas were likely a mosaic of forested and open woodlands

providing ideal habitat for large herbivores such as horse, red deer, wild boar and aurochs.

In the inland limestone Massif cave sites, Picareiro and Caldeirão, red deer, caprids and

rabbit were the main prey species (Bicho 1994b; Zilhão 1992). These are thought to represent

short-term logistical extraction sites used by small groups of hunters who made forays

from residential camps in the valleys, now mostly buried under Holocene alluvium and


An increase in the number of sites from 20 in the first phase (an average of 3/1000

years) to almost 40 in the second phase (an average of 16/1000 years) characterizes the

second settlement phase. Additionally, there was an increase in the number of primary

areas of occupation used by the Epipaleolithic hunter-gatherers, including coastal areas

to the south of the Tagus and Sado Rivers in Alentejo. The types of sites found during this

period include the shell-bearing inland caves and rockshelters, coastal shell middens, and

open-air sites on the flat plains of the interior river valleys (Bicho and Haws 1996). The

maritime resources brought to the cave/rockshelter sites show clear evidence for trips to

and from the coast.

Expansion into previously unoccupied Alentejo by Epipaleolithic hunter-gatherers

suggested a rise in regional population. The sharp change in settlement system was argued

to be a result of demographic expansion similar to northern Spain (Straus and Clark 1986).

However, ‘expansion’ may not be the proper term. As Zilhão (1990) has pointed out, the

estimated 12,000 km2 area of Estremadura during the Solutrean was enough to support a

maximum band of almost 500 people if one accepts the proposed Upper Paleolithic

population density of 0.04 people/km2 (Cohen 1977; Wobst 1974). If this were the case,

then rising sea level in the Late Pleistocene may have had a significant impact on human

groups by shrinking the size of available land area by 25% to 8,700 km2 (Perlman 1980).

This, in turn, may explain the expansion of human occupation out of Estremadura into

Alentejo during the Epipaleolithic. Demographic expansion has also been put forth as an

explanation for the Mesolithic land use and settlement system (Straus et al. 1990). It is

unclear whether there were periods of punctuated population growth or a more gradual

trend of increasing population, that led to changes in settlement patterns. In fact, there

may never have been any significant increases in population (Hayden 1981a). The increases

in numbers of sites may have resulted from increasing logistical settlement patterns, which

would have simply created more types of sites (Binford 1980; Straus 1991c). It may also be

that the sites from earlier periods are simply buried in marine sediments off the coast or

were destroyed by wave action during the sea level rise. Until more sites are found through

systematic survey and more is known about the effect regional geomorphological changes

may have had on site preservation, these ideas remain to be tested.



The Cantabrian and Mediterranean cases have important implications for explaining

Late Upper Paleolithic adaptations in Portugal. The models for central Portugal discussed

above point to a similar trend of increasing specialization and diversification in central

Portugal (Zilhão 1992; Bicho 1993, 1994; Bicho and Haws, 1996). But when the Cantabrian

data are considered several questions become apparent. First, does the presence of marine

fauna and small game in the Late Upper Paleolithic of central Portugal mean that

populations were stressed and forced to broaden their niche? Alternatively, was it based

on choices made to exploit reliable resources and thus reduce risk as was implied for

Mediterranean Spain?

A second problem concerns plant consumption during the Upper Paleolithic in central

Portugal. The excavation of El Juyo in Cantabria and Nerja in Andalucía has shed some

light on this problem in Spain (Freeman et al. 1988; Badal 1998). However, the lack of data

has led to only brief remarks acknowledging the likelihood of plant consumption in central

Portugal (Bicho 1993; Zilhão 1997). No attempts have been made to understand the factors

involved in plant exploitation which, in the absence of substantial direct evidence, ought

to begin with an understanding of human nutritional requirements, ethnographic

observations of recent hunter-gatherers, consideration of plant availability through

paleoenvironmental reconstruction and information on the nutritional benefits and

productive yields of edible wild plants.

Third, models of intensification and specialization were based on the percentages of


various animal taxa in each site and preliminary indications from faunal analyses of key

sites such as Caldeirão and Picareiro. Detailed faunal analyses of assemblages from

Picareiro and Suão caves are presented in the next section for the first time. These data are

compared to the record from Mediterranean Spain where similar environmental conditions

and human adaptations occurred during the Late Pleistocene and Early Holocene.

Afterwards, the previous models are evaluated using the results presented here.

Part II: Prehistoric diet and subsistence

Reconstructing past subsistence necessitates an understanding of the availability of

edible plants and animals and a means of estimating the proportions of these incorporated

into the diet. The former is usually accomplished by paleoecological studies, archaeofaunal

and archaeobotanical studies. The latter usually relies on ethnographic and experimental

data and theoretical models such as those discussed in Chapter 2. Essentially, archaeologists

have two major means of studying prehistoric diets: (1) archaeological remains, in this

case archaeofaunal and floral remains, and (2) human bone chemistry, mainly stable

isotopes and trace elements. Indirectly, paleoenvironmental reconstructions based on

pollen diagrams, charcoal analysis, micromammal assemblages, deep sea foramininfera

and general circulation models can enable resource availability modeling.

The ‘Man the Hunter’ conference and volume revolutionized archaeologists’

perceptions of hunter-gatherers. It destroyed the long held assumption dating back to

Hobbes that hunter-gatherers worked exceptionally hard to eke out a living on wild plants

and animals. Lee and DeVore (1968) were successful in demonstrating how well hunter-

gatherers lived even in the most marginal environments on the planet. Even though it is

widely cited, to the point that almost every paper on hunter-gatherers begins with a

statement, “…ever since ‘Man the Hunter’…,” many of the important ideas from the volume

have been ignored or reinterpreted. Perhaps the main reason behind this is the adoption

of foraging theory in ethnographic and archaeological studies. The “affluent” San, who

spent very little time in the food quest are now “marginal” or “suboptimal” because they

are forced to subsist off of “low-ranked”plants for the majority of their diet with the

occasional large game animal. Hawkes and O’Connell (1985) “exposed” the mongongo

nut dilemma by pointing out the high processing costs involved. However, even with

high processing costs offsetting the caloric gain (thus resulting in lower return rates) the

fact remains that the San still do not spend much time gathering and processing food.

There must be some other reason behind this. As foraging theory is applied to

archaeological cases, large game animals are generally considered the highest ranked

Logical arguments suggest and empirical data from experimental and
ethnographic settings demonstrate (see Broughton 1994b for references and
discussion) that, for singly-handled animal prey, post-encounter return rates
are generally scaled to prey body mass. Among Holocene North American
vertebrates in particular, the larger the animal, the higher the post-encounter
return rate. This fact, combined with the proposition that overall foraging
return rates declined in the late Holocene, leads to the prediction that low-
ranked (smaller-sized) vertebrates should have become more important in
human diets at this time... (Broughton and O’Connell 1999: 155).
Accordingly, if people are subsisting only occasionally on large game and mostly

plants or small animals, then they are acting suboptimally or living in a poor environment.

This marginalization is due primarily to over-harvesting of high-ranked game, also known

as resource depression, or possibly some other feature that has lowered the return rate for

the highest ranked resource, thus forcing people to add a previously uneaten, lower ranked

resource. Theorists consistently make the point that no matter how abundant a resource

becomes a forager will not adopt it unless a higher ranked resource no longer provides

the high return rate. Therefore, regions with generalized hunter-gatherer diets probably

reflect low frequencies of highly ranked resources, most often large game. Either the

environment is ‘poor’ to begin with or resource depression has occurred due to climate

change or over-harvesting.

Broadly speaking, archaeologists studying Paleolithic and Mesolithic subsistence in

Europe take for granted specialized large game hunting economies in the former and the

generalized Broad Spectrum Adaptation in the latter. This is due to a number of reasons

such as a biased archaeological record of animal bones taken as an accurate reflection of

past human diet, theoretical emphasis on energy-based foraging models and the intensity

of research in areas where glacial climatic conditions curtailed plant productivity. There

is an underlying assumption that holds human diet became much more meat-focused

early in human evolution arguably related to brain evolution and colonization of temperate

Eurasian latitudes (Isaac and Crader 1981; Milton 1987; Aiello and Wheeler 1995; Rolland

1998; Stiner 2002). This trend culminated in the hunting economies of Middle and Upper

Paleolithic peoples in Eurasia. Plants, aquatic resources and small game entered the diet

only after large game populations became depressed due to hunting pressures brought

on by ever-increasing Pleistocene human populations. Recent stable isotope and trace

element studies lend support to the idea that Neandertals were highly carnivorous while

early modern humans ate a slightly more diverse diet (Bocherens et al. 1991, 1995, 1999;

Richards et al. 2000a&b, 2001). The explanations for this difference include population

pressure forcing humans to expand their diets (Stiner 2001), niche differences between

two competing human populations (Smith 2002), and nutritional decisions resulting in

greater reproductive success (Hockett and Haws, in press).

Southern Europe has been called a “refugium” for temperate-loving plants and animals

by many authors (e.g., Straus 1991). Because of the focus on the Franco-Cantabrian region,

researchers often follow the explanatory framework utilized for northern Europe (Straus

1996). Ironically, the Broad Spectrum Revolution of the northern European Mesolithic

occurred at the same time the climate improved, whereas it occurred earlier in southern

Europe and the Near East where Late Pleistocene climate was much less severe and the

scale of environmental change less dramatic. This is argued to reflect differences in the

timing of population pulses (Stiner 2001). However, low population densities were not

the reason Upper Paleolithic sites in northern Europe lack hazel nutshells. Hazel shrubs

simply were not present. In the south, especially Iberia, temperate and Mediterranean

plant and animal species were always present but in differing proportions.

Unfortunately, Upper Paleolithic people in Portugal did not bury their dead in geologic

contexts suitable to preservation making stable isotope and trace element studies

impossible. With the exception of a few isolated teeth and skeletal fragments from about

four sites, the only intact burial is that of the alleged hybrid child from Lagar Velho (Duarte

et al. 1999). Since this skeleton is of a four year old child it may not be representative of the

“average” person’s diet. That said, the nearby and possibly contemporaneous (at least in

gross terms), hearth contains abundant ungulate and even marine mammal (dolphin)

remains which provide glimpse of a diverse early Upper Paleolithic diet and coastal/

inland movement pattern.

Given the lack of human remains, the only current approach to the direct study of

Upper Paleolithic subsistence in Portugal is the archaeofaunal and botanical record. The

record is biased entirely in favor of animal bones, so inferences about plant use can only

be made by generating hypotheses using indirect sources of palynological and

anthracological (charcoal) data, ethnographic observations of “modern” hunter-gatherers,

foraging theory and human nutritional ecology. Taxonomic lists of fauna from

archaeological and paleontological sites and plant inventories from pollen diagrams and

charcoal analyses can provide a potential range of available food resources to prehistoric

people. Seasonality studies do not simply tell when a site was occupied, but permit models

of seasonal mobility when combined with resource availability estimates and nutritional

yield data. Detailed taphonomic studies of faunal assemblages provide more accurate

pictures of resource use and site function. Together these approaches allow a better

understanding of subsistence and settlement patterns than those based solely on

technological organization and site location.

In the following chapters, paleoenvironmental data and the regional archaeological

record are used to evaluate plant resource use through the lenses of taphonomy, foraging

theory and nutritional ecology. Data on animal resource use from the faunal-bearing sites

in Estremadura are then used to understand the nature of large and small game utilization,

site function, and temporal trends from the Late Pleistocene and Early Holocene.

Chapter 4: A Western Mediterranean perspective on Upper Paleolithic plant


4.1 Plant use by prehistoric Mediterranean hunter-gatherers

David Clarke, in his essay “Mesolithic Europe: The Economic Basis” (1976), challenged

the “meat fixation” of archaeologists and highlighted the potential for intensive plant use

in Europe prior to the introduction of agriculture. He argued that plants likely made up

60-80% of the diet of prehistoric hunter-gatherers in Europe. Indeed, ethnographically

known hunter-gatherers in temperate regions incorporate such percentages of plants in

their diet (Kelly 1995). The only area where meat completely dominates is the Arctic,

where the lack of edible plants during much of the year leaves little choice. Even in this

extreme, people are known to eat the contents of reindeer stomachs and exchange meat

for seaweed with coastal peoples (Clark 1952). Even still, this may only reflect a seasonal

aspect of subsistence. The notion that meat would comprise 90% of the diet during the

Upper Paleolithic or Mesolithic seems unfounded given these facts about modern hunter-

gatherers and the nutritional problems associated with excess protein consumption. Recent

stable isotope analyses suggest such a meat-focused diet in Britain during the LGM

(Richards et al. 2000), although one could argue that this is merely analogous to the Arctic

case and is not the “rule” for all of Europe during this period. Much of Iberia, especially

Mediterranean Spain and Portugal, never experienced dramatically cold extremes except

in mountain zones. Therefore, it seems reasonable to assume a high degree of plant

exploitation in this part of Europe at least as far back as the LGM and likely farther. Even

at the Cantabrian Magdalenian site El Juyo, where climatic and environmental conditions

were cooler and less-vegetated, Freeman et al. (1988) reported plant macrofossil remains

from 21 families and 51 genera. These included acorn and hazelnut fragments, berry pits,

grass seeds and aquatic plants. During this same period, prehistoric Mediterranean hunter-

gatherers would have had access to a wide variety of plants including oak acorns, hazelnuts,

wild tree fruits, berries, edible bulbs, grass seeds, and legumes.

Interestingly, the potential for plant exploitation during the Upper Paleolithic has

never received due attention (Mason et al. 1994). The focus has always been on the animal

remains from archaeological sites. Most authors discussing Paleolithic subsistence either

do not mention plants or suggest they would have been eaten when available and then

dismiss the significance of their contribution to prehistoric diet and subsistence economies.

Many researchers have argued that plants were not regularly exploited because the Late

Pleistocene environments of Europe were too extreme to permit stable plant communities.

Richerson et al. (2001) have recently suggested that low global CO2 levels during the Late

Pleistocene kept plant productivity suppressed thus making plants unattractive to human

foragers. They further argued that plant-based diets would have taken a long time to

develop and that frequent climatic oscillations made long-term adaptations to plant

resources unlikely. Stiner and Kuhn (2001) have also argued that intensive plant

exploitation is only visible archaeologically at the end of the Pleistocene when stone

processing technology appears, namely grinding stones. This echoes Hayden’s (1981)

earlier claims.

Clarke’s (1976) model for the Mesolithic might have been more acceptable to

archaeologists working in that period because the Early Holocene was precisely the time

that the Broad Spectrum Revolution was said to have occurred. For the Upper Paleolithic,

the ‘meat fixation’ was unchallenged. The main obstacles are the acceptance of energy as

the appropriate currency to measure human subsistence choices and the invisibility of

plants and the technology used to process them in the archaeological record. As discussed

in chapter 2, Kelly (1995) and others have argued that animal resources are usually better

sources of energy than plants. In general, plants are viewed as low-ranked resources

because they often have high percentages of inedible cellulose and/or toxins that must be

removed prior to human consumption. The high processing costs outweigh their ease of

collection and natural abundance when caloric value is relatively low. Since most foraging

models consider all foragers equal, plants usually rank low. Some argue that any nutrient

value will be greatly diminished or lost entirely during processing. However, as Stahl

(1989) has pointed out, processing plants does not necessarily mean there will be nutrient

loss. In many cases, toxins must be removed, thus increasing the nutritional value. Cooking

may increase certain nutrient values in plants (Wandsnider 1997; Wrangham et al. 1999).

In some cases, the processing costs may be offset by the benefits of nutrient increase

although there is only qualitative data to support this because foraging modelers equate

processing with nutrient loss (Stahl 1989).

Many authors have assumed that extensive and intensive plant exploitation began

with the appearance of grinding stone technology widely used to process seeds and nuts

(Hayden 1981; Kuhn and Stiner 2001). However, recent reports show unaltered stone

tools are used by chimpanzees in Africa to crack open nuts (Mercader et al. 2002). The

Lower Paleolithic site of Gesher Benot Ya’aqov has nut-cracking stones associated with

several species of nuts (Goren-Inbar et al. 2002). Thus, stone tools have been used to process

plants for a considerable time. Proponents would likely counter that these are simple

tools which require minimal labor input and that intensive plant processing, especially

seeds, was only made possible when grinding stones were invented. Still, this should not

be considered a requisite for intensive plant exploitation. The fact is that many recent

hunter-gatherers used perishable wooden implements to pound seeds and nuts (Lee 1979;

Stahl 1989). Even stone sickle blades are not necessary to harvest wild grains, as pointed

out by Madella et al. (2002) in their study of plant exploitation at the Middle Paleolithic

site of Amud in Israel.

Another problem with plant visibility is the lack of preservation. Mason et al. (1994)

have called for greater attention to recovering plant remains from Paleolithic sites. The

main problem is that most plant foods would leave little trace in many sites. Preserved

plant food remains have never been found in Upper Paleolithic sites in Portugal, but they

have been recovered in archaeological sites around the Mediterranean. Numerous charred

pine nut cones and shells were found in Middle Paleolithic hearths at Vanguard and

Gorham’s Caves in Gibraltar (Gale and Carruthers 2000) and throughout the Upper

Paleolithic sequence at Cueva de Nerja (Badal 1998, 2001). Magdalenian and Epipaleolithic

levels of Nerja also contained acorns and wild olive pits (Aura et al. 1998). Also in Spain,

Freeman (1981) reported charred grass and vetch seeds from Abric Agut, which has now

been radiometrically dated to the Pleistocene-Holocene transition (Vaquero et al. 2002). In

Greece, Koumouzelis et al. (2002) found evidence for seed-bearing grasses including

Chenopodium and Polygonum, Silene and other typical Mediterranean plants in the

Aurignacian site, Klisoura Cave. Plant foods were recovered from Upper Paleolithic levels

at Franchti Cave (Hansen 1978). At Ohalo II in Israel, Kislev et al. (1992) reported acorns

and several other plants dated to 19,000 bp.

Evidence for the consumption of hazelnut, wild fruit and several legumes in the Early

Holocene has been found at Balma Arbeurador in France, dated to 8,740 bp (Vaquer et al.

1986). In Catalunya at Cingle Vermell (dated 9,760 bp) numerous remains of hazelnuts,

acorns, pine nuts, chestnuts and wild fruits have been recovered (Vila i Mitja, 1985). Nearby,

at Roc del Migdia (7,000-9,000 bp), Holden et al. (1995) have identified hazel shell, sloe and

parenchymatous tissue of roots, tubers and edible aquatic plants. In the Basque country

of Spain, Peña (2000) has reported on Mesolithic and Neolithic wild plant gathering. In

several sites hazelnuts, acorns and wild tree fruits were common in both periods. Olària

et al. (1988) reported pine nuts, acorns and chestnuts at Cova Fosca (9,460 bp). At the

Mesolithic site, Grotta dell’Uzzo, in Sicily wild peas, acorns and arbutus fruits were

recovered (Costantini 1989). These examples illustrate the widespread prehistoric use

and richness of edible plant resources in the Mediterranean region. The rest of this chapter

focuses on the Iberian Peninsula with primary emphasis on Portugal. Again, the central

region of Estremadura is the center of attention as this area of Portugal is the best known.

4.2 Plant resources in Iberia

Present-day climate and vegetation in Portugal

Several forces determine the climate of Portugal and create its mixed Atlantic and

Mediterranean environment. In the winter, the Polar Jet descends into the North Atlantic

and brings storms across the ocean to Iberia. In the summer, the Azores High pressure

cell keeps moisture from penetrating southward, thus creating the summer dry season.

Annual rainfall varies from 1000-2000 mm in northern Portugal to 300 mm in the south.

The central coastal regions average about 600-700 mm annually. In most of the country,

the majority of the precipitation falls between October and March. Average winter

temperatures range from 8-12° C on the northern coast to about 16° C in the south. The

interior north and montane areas temperatures often fall below freezing in winter. Summer

temperatures range from average highs of 25-28° C near the coast to as high as 40° C in the

interior. In Estremadura, winters are cool and wet while summers are warm and dry.

On the Iberian Peninsula, the Euro-Siberian vegetation typical of temperate Europe

occurs along the northern strip from the Basque country in the northeast to northern tip of

Portugal in the west. The rest of the peninsula is covered by Mediterranean-type vegetation.

Rivas-Martinez (1982) divided the Mediterranean vegetation into étages or levels based on

altitudinal zonation. Most of Portugal falls within the meso-Mediterranean zone, which

in its natural state is characterized by evergreen oak forest. The typical species are listed

in Table 4.2. In central Portugal, meso-Mediterranean tree and shrub communities are

found in varying proportions depending on exposure, soils and altitude. Agriculture,

intentional planting and timber harvesting over several millennia have dramatically altered

the natural vegetation communities. No truly wild places exist any longer. Some areas

have been protected for several centuries and their composition provides the best model

for the natural distribution of trees and shrubs. In these places, the evergreen oaks, Stone

Table : Mediterranean bioclimatic altitudinal zones

Bioclimatic étages Average annual Average low Average high Thermic Index
temp winter temp winter temp
Altimediterranean ° C 
° C ° C   
Oromediterranean  ° C 
 ° C  ° C   

Supramediterranean   ° C  ° C  ° C  

° C  ° C  ° C  

 ° C  ° C   ° C  

After RivasMartinez (

) cited in Vernet (


Mediterranean Sea

  km  SupraMediterranean

Figure : Map of bioclimatic zones in Iberia


Table  : Some presentday trees and shrubs native to Portugal

SupraMediterranean Latin name English name Portuguese name
trees Quercus pyrenaica Pyrenean oak carvalhonegral
Q robur common oak carvalhoroble
Q faginea Portuguese oak carvalhocerquinho
Pinus pinaster Maritime pine pinheirobravo
Castanea sativa chestnut castanheiro
Sorbus aucuparia Rowan ash tramazeira
Alnus glutinosa alder amieiro
Prunus avium wild cherry cerejeirabrava
Betula celtiberica birch vidoeiro
wetland trees Ulmus minor elm ulmeiro
Fraxinus angustifolia ash freixo
Salix sp willow salgueiro
Populus alba poplar

shrubs Juniperus communis juniper zimbro

Taxus baccata yew teixo
Prunus spinosa sloe
Corylus avellana hazel aveiro
Ilex aquifolium holly azevinho
Cytisus sp broom
Erica sp heath urze
Genista sp piornais
trees Quercus ilex Holm oak azinheira
Q suber cork oak sobreiro
Q coccifera Kermes oak carrasco
Q faginea Portuguese oak carvalhocerquinho
Pinus pinaster Maritime pine pinheirobravo
P pinea Stone pine pinheiromanso
Phillyrea latifolia privet pelo aderno
Acer monspessulanum Montpellier maple zelha
Arbutus unedo wild strawberry medronheiro
Olea europaea var wild olive zambujeiro
Buxus sempervirens box
Acer pseudoplatanus sycamore bordo
Laurus nobilis laurel loureiro
wetland trees Salix sp willow salgueiro
Fraxinus angustifolia ash freixo
Populus alba poplar choupo
Ulmus sp elm ulmeiro
Sambucus nigra elderberry
shrubs Juniper oxycedrus prickly juniper cedro espanhol
Erica arborea tree heath urze
Myrtus communis myrtle murta
Rhamnus alaternus Mediterranean sanguinhodassebes
Pistacia lentiscus mastic aroeira
Jasminum fruticans wild jasmine jasmineirodomonte
Asparagus albus asparagus estrepes
Rosmarinus officinalis rosemary alecrim
Viburnum tinis laurustinus folhado

pine, privet, wild strawberry, wild olive, myrtle and Mediterranean buckthorn prefer the

sunny, exposed areas. The Portuguese oak, Montpellier maple, laurel and Maritime pine

are found in shadier areas. Along streams other trees and shrubs include alder, elm,

poplar, willow and elderberry. The deciduous oak, Quercus faginea, is often seen as a

transitional species occurring in the north and south (Vieira et al. 2000). In the north and

in the mountains, the vegetation is predominately supra-Mediterranean. Arboreal

vegetation includes the deciduous oaks, chestnut, pine, birch, ash, elm, willow, and wild

fruit trees. Shrubs include juniper, yew, hazel, holly and heather.

In discussing southern Europe, Clarke (1976) wrote that the Mediterranean woodlands

offered 200-350 edible wild plant species. He suggested numerous wild grass seeds, roots,

bulbs, herbs and legumes were widely distributed and abundant on the Early Holocene

Mediterranean landscape. Examples of these native to the Iberian Peninsula are shown in

Table 4.3. Combined with fruits, berries and nuts, hunter-gatherers would have maintained

a diverse and nutritious diet when these plant foods were consumed with animal resources.

Pine nuts in Iberia

In particular, Clarke (1976) focused on the high protein, high yield seed of Pinus pinea,

the Stone pine. Pine nuts are well known as a food source in Europe. The Greeks considered

them (and acorns) ‘food of the gods’ (Howes 1948). Abundant remains are also known

from Roman camps in northern Europe, outside the ecological range of the species. As

noted above, their charred shells are found in Middle Paleolithic and Upper Paleolithic

contexts. Yet, despite their early use, pine nuts (and most other mast resources) have been

Table  : A sample of edible wild plants on the Iberian Peninsula

Latin name English name edible part nutritional value

Cruciferae Sisymbrium officinale hedge mustard leaves essential oils

Cardaria draba hoary cress leaves

Caryophyllaceae Silene vulgaris bladder leaves shoots vitamins
Stellaria media chickweed dried flowers medicinal vitamins
Portulacaceae Portulaca oleracea purslane leaves vitamins fatty acids
Chenopodiaceae Chenopdium sp goosefoot/ fat leaves seeds vitamins
Beta maritima wild beet leaves
Salicornia europaea glasswort entire plant vitamins minerals

Malvaceae Malva sylvestris common leaves essential oils

Rosaceae Rubus sp blackberry leaves berries Vitamin C citric acid
Sorbus aucuparia Rowan ash berries Vitamin C
Prunus spinosa sloe berries sugars

Umbelliferae Coriandrum sativum coriander fruit seed

Apium sp wild celery leaves
Petroselinum crispum parsley leaves stems Vitamin C
Carum sp caraway leaves roots
Foeniculum vulgare fennel rhizome leaves starch fatty acids
Polygonaceae Polygonum sp knotgrass seeds
Rumex crispus curly dock leaves Vitamin C

Urticaceae Parietaria diffusa pellitoryof dried flowers medicinal

Ericaceae Calluna vulgaris heather dried flowers medicinal

Verbenaceae Verbena officinalis vervain dried flowers medicinal

Labiatae Salvia officinalis sage leaves essential oils

Rosmarinus officinalis rosemary leaves essential oils
Lavandula angustifolia lavender dried flowers essential oils

Plantaginaceae Plantago major plantain leaves Vitamin C

Caprifoliaceae Sambucus nigra elderberry fruit essential oils sugar

Compositae Sonchus oleraceus sowthistle leaves Vitamin C

Liliaceae Allium sp wild onion bulbs leaves Vitamin C

leek shallot
Typhaceae Typha latifolia cattail rhizome young starch
shoots pollen
Phycophyta Phaeophyta seaweed blades or Vitamin C
Chlorophyta seaweed blades or vitamins minerals
Rhodophyta seaweed blades or vitamins minerals
Information from Guil et al ; Launert  

marginalized by archaeologists because they are considered an unpredictable and therefore

unreliable resource.

In western North America, much has been written on the subject since piñon nuts

were known in the subsistence of many Great Basin groups. Their widespread distribution

and abundance made pine nuts an important, and probably high-ranked food item in the

prehistoric diets of native peoples (Steward 1938; Thomas 1973). Simms (1987) calculated

a relatively low kcal/hr return rate for pine nut processing in the Great Basin. However,

among plants they ranked high and Simms predicted that they would be collected before

most other resources. Because the masts occur every 3-5 years, hunter-gatherers would

need to constantly monitor stands. Recently, Sullivan (1992), in challenging the maize-

centered diet for the Western Anasazi of the Grand Canyon area, postulated that prehistoric

hunter-gatherers with ample knowledge of the land in which they lived would have been

able to predict mast years and/or move to areas where piñon harvests would have been

abundant. He interpreted fire-cracked rock piles in a number of sites as evidence of piñon

roasting, although paleobotanical remains were lacking. In a recent paper, Mithen et al.

(2001) have interpreted similar features in Colonsay off Scotland as hazelnut roasting ovens.

No substantial features such as these exist in the Mesolithic or Paleolithic archaeological

record in Portugal, but these are not prerequisites as the charred shells from Gibraltar and

Cueva de Nerja show.

Several species of pines are found naturally on the Iberian Peninsula today. These

include Pinus pinaster and P. pinea in Portugal. In Spain these two exist along with Pinus

nigra, P. halepensis, P. uncinata and P. sylvestris. Because of widespread intentional planting


of P. pinaster and P. pinea for centuries and probably even millennia, it is virtually impossible

to know their natural ranges. It is thought that both evolved on the Iberian Peninsula

(LeMaitre 1998). Because the seed of P. pinea is the commonly eaten one, the focus will be

on this pine. However, this does not necessarily mean that P. pinaster seeds would not

have been exploited. Rhode and Madsen (1998) noted that the much smaller limber pine

seeds were utilized prior to piñon in the Great Basin.

Pinus pinea lives in sandy soils and podzols in sunny areas of Iberia up to 1,000 m asl.

This species does not thrive in highly calcareous soils but can tolerate a limit of 50%

limestone (Vieira et al. 2000). It is generally found in thermo- and meso-Mediterranean

regions and thrive in average annual temperatures 13-19° C. Their low temperature

tolerance is about -5° C. They can withstand annual rainfall as low as 250-300 mm but

prefer 400-1,000 mm. This species masts every 3-4 or 5-6 years but some seeds are produced

during non-mast years. In stands in southern Portugal, trees typically produce 250 cones

per year, but often 1,000 and sometimes 2,000 (Vieira et al. 2000). Seed production is

generally more reliable in P. pinea than pines of western North America (LeMaitre 1998).

Peak production occurs in November and cones remain on the trees until January or

February. Each cone averages about 100 pine nuts which is a tremendous contrast with P.

monophylla of western North America that typically produces 6-16 nuts per cone (Barlow

and Metcalfe 1996; Badal 1998). Similar yields were observed in natural stands of P. pinaster

(Miguel Pérez et al. 2002). Therefore, a single tree could produce 25,000-100,000 and even

200,000 pine nuts per year. In central Spain, where annual rainfall averages 450 mm,

natural pine forests contain about 100 trees per hectare (Miguel Pérez et al. 2002). Similar

conditions in the past would have presented prehistoric hunter-gatherers with a bountiful

harvest. Modern workers harvesting P. pinea with long pikes collect 400-600 cones per

day on average. Cones are left exposed to the sun to dry and open so that seeds can be

collected. One hundred kg of cones yields 15-22 kg of hulled seeds. For pine nuts in their

hulls, 1 kg equals about 1,400 seeds. For hulled seeds, 6,300 pine nuts weigh about 1 kg

(Badal 1998). With minimal time and labor investment, prehistoric hunter-gatherers could

have easily harvested and processed substantial amounts of pine nuts.

Table : Comparison of nutritional values () of different types of

pine nuts
Protein Fat Carbohydrate kcal/g
Old Worl d
Pinus pinea     
P sibirica  
P gerardiana   
Ne w World
Pinus monophylla  
P edulis   
P cembroides    
P quadrifolia 
P sabiniana    
P strobiformis 

Data from Howes ( ) Barlow and Metcalfe () and Lanner
( )

Table 4.4 shows the nutritional values for some of the varieties of pine nuts utilized

for food around the world. There is considerable variability in pine nut nutrient

composition between species. The most striking characteristic of P. pinea is the high

percentage of protein. It also has high percent fat but low percent carbohydrate. In contrast,

the values for P. monophylla in the Great Basin show it is poor in protein but high in

carbohydrate. Considering Speth’s (1991) argument that hunter-gatherers could have


avoided problems associated with excessive protein intake by consuming carbohydrates,

P. monophylla may have been a valuable ‘protein-sparing’ resource in the Great Basin.

However, in Iberia, pine nuts may have been more important as storable reserves of

balanced protein and fat in order to mitigate the drought stress of summer. Table 4.5

shows nutrient values for P. pinea measured in grams/100g by three different laboratories.

These results show that despite a high protein percentage, P. pinea seeds do constitute

good sources of carbohydrate.

Table : Comparative nutritional values expressed in g/g for selected nuts
from Mediterranean woodlands
Protein Fat Carbohydrate kcal
Pinus pinea    
Q suber  
Q ilex     
Castanea sativa  
Juglans regia     
Corylus avellana  

Data from USDA food composition database; Alce Ingeniería (Spain)  Tabela da
Composição dos Alimentos Portugueses (Gonçalves Ferreira & da Silva Graça

Seeds of P. pinea have a high fat content and therefore need to be dried or roasted to

avoid spoilage. In controlled conditions, seeds dried at room temperature showed increases

in soluble protein content that peaked after 6 months (Fernández-García de Castro and

Martínez-Honduvilla 1982). Under low heat (47o C) seeds showed an overall increase in

protein and free amino acids but a slight decrease in lipid content (Martínez-Honduvilla

et al. 1974). Although no data were presented on changes in fat content in stored seeds it

may be reasonable to assume that low-heat drying would result in better fat-retention. In

addition, pine nuts can be eaten raw but their flavor and storability improves with light


Acorns in Iberia

For almost a century, anthropologists have realized the importance of acorns as human

food (Merriam 1918). Anthropologists long held that the incredible natural abundance of

different acorns attracted human groups and led to their role as a dietary staple. Acorns

seem to have supported very large human populations in late-prehistoric and early historic

California. Until recently, acorns were viewed as a highly nutritious and productive

resource that could be easily collected and utilized. Basgall (1987), using principles from

optimal foraging theory, noted that acorns actually required significant labor investment

to leach the tannins that made them too bitter for immediate consumption. He argued

that the caloric return rates for acorns made them a low-ranked item because of the high

processing costs relative to energy yield. Among North American archaeologists, acorns

quickly became perceived as a marginal resource that only became economically important

when other resources were exhausted (Basgall 1987; Mason 1995a&b). However, this model

is based on the assumption that the appearance of grinding stones in California prehistory

was correlated with acorn consumption (Basgall 1987). Since acorns can be intensively

processed using perishable materials such as wooden mortars and pestles, earlier acorn

use might be archaeologically invisible. Additionally, evidence of leaching methods such

as soaking in water either in special pits or in streams, boiling and roasting might not

survive. Therefore, dismissal of significant prehistoric acorn consumption may be



In his book, Prehistoric Europe: The Economic Basis (1952), Grahame Clark discussed

prehistoric acorn consumption in the Mediterranean. Citing the geographer, Strabo, he

noted the Lusitanians, in what is now Portugal, were observed to eat bread made of ground

acorns for three-quarters of the year. Although in later times acorn flour was milled and

made into “famine breads” when grains were scarce, many people appear to have subsisted

off acorns for centuries (Jørgensen 1977). Numerous citations from classical sources suggest

acorns were viewed as the basis for all of civilization (Clark 1952; Mason 1995; Vencl 1996;

Sieso and Gómez 2002). In fact, the genus name “Quercus” is derived from two Celtic

words meaning “beautiful tree” suggesting its importance in early times (Sánchez Arroyo

1999). Vencl (1996) and Sieso and Gómez (2002) provide detailed summaries of the

archaeological evidence for acorn use through time on the Iberian Peninsula. In Portugal,

Afonso do Paço (1954) reported concentrations of acorns at the Copper Age site Vila Nova

de São Pedro. Senna-Martinez (1994) reported carbonized acorns from the Neolithic site

of Ameal in northern Portugal. Bicho (1993) and Arnaud (1990) hypothesized that acorns

were eaten during the Late Upper Paleolithic and Mesolithic though no direct evidence

has been recovered to date. Acorn-eating, or balanophagy, survives today in Iberia where

sweets are made from acorns. On Sardinia, local people still gather acorns and process

them using traditional methods. Acorns are mixed with a special iron-rich clay and boiled

to absorb the tannins (Johns 1998). In the western Rif of Morocco, acorns are eaten raw,

toasted, soaked in water or sun-dried (Peña 2000).

Mason (1995b) detailed the widespread use across space and time in her review of

the use of acorns as food in prehistoric Europe. She gave four reasons why acorns may

have played an important role in Mesolithic diets which are also applicable to the

Mediterranean Paleolithic and Epipaleolithic:

1) Oaks were an important element of vegetation…an would have been a
commonly, and sometimes abundantly available, resource.
2) Acorns are, in nutritional terms, a potential energy staple— they are a
bulk carbohydrate-provider and are very similar in nutritional terms to the
3) They are potentially storable for long periods, and can be a valuable
overwintering resource.
4) Ethnographic and documentary evidence from wherever oak trees are
found suggests that acorns have been an important (often dominant) plant-
food resource for hunter-gatherers…(2000: 141).

Prior to the adoption of cereals, acorns may have been significant resources for

prehistoric Mediterranean hunter-gatherers as is well-documented for Eastern and Western

North America and Japan (Merriam 1918; Lewthwaite 1982; Mason 1995a&b). In the Near

East there is solid evidence that acorns were used as food as early as 19,000 bp at Ohalo II

(Kislev et al. 1992). At La Sarga, an Epipaleolithic site in València, a painted rock art scene

shows several figures collecting acorns as they fall from the tree (Fortea and Aura 1987).

However, inadequate recovery techniques and/or preservation biases inhibit an

understanding of the role acorns may have played in European hunter-gatherer subsistence.

The absence of acorn parts is made difficult by taphonomic factors such as possible small

mammal or wild boar feeding (Leiva and Fernández Alés 2003). Processing methods may

also disfavor preservation as most remains may never have been charred and thus

preserved more easily (Mason 1995). Field processing among stands near base camps

may also eliminate those parts of the acorn (or pine nut) so that the evidence for use is

discarded offsite. These processing locales would likely be archaeologically invisible since

they would not require the use and retouching of stone tools which would leave behind a

durable residue. If, however, processing included roasting and/or pounding with stones

a durable record might be left. In this case it is doubtful, in the absence of lithic type

fossils, that such scatters of unflaked, unretouched stones and charcoal would be reported

as an Upper Paleolithic site.

Iberia is home to several species of evergreen and deciduous oaks. Typically, evergreen

oaks are found in the thermo- and meso-Mediterranean vegetation zones. These include

Quercus ilex, Q. suber and Q. coccifera. Deciduous oaks are found in the meso-Mediterranean

zone but mainly in the supra-Mediterranean. These species include Quercus faginea, Q.

pyrenaica, Q. robur. The species most commonly used as food in Iberia throughout the

past is Q. ilex, but each can be consumed with some degree of processing. Q. ilex and Q.

suber produce sweet acorns that require minimal processing time, usually light roasting.

Q. coccifera has an astringent and bitter acorn and would require considerable leaching.

The deciduous oaks are slightly bitter compared to Q. ilex and Q. suber, however they

were consumed by people in the past (Mason 1995; Peña 2000; Sieso and Gómez 2002).

The Holm oak, Quercus ilex, today prefers calcareous soils and rainfall between 600-

1000mm. Holm oaks cannot maintain closed canopies with less than 400mm precipitation

(Terradas 1999). Below that, Mediterranean shrubs will dominate. Q. ilex is more cold-

resistant than other evergreen oaks in Portugal. Today it withstands temperatures as low

as -15° C and can live through short periods as low as –20 to -25° C. Young trees may not

survive below -10° C so that may be the low point for overall survival (Terradas 1999).

The Holm oak is a mast species in which high acorn production occurs every 4-6 years.

Acorns mature and fall between November and January. Data on Holm oak productivity

in Catalunya show considerably inter-annual variability in acorn production. Siscart et al.

(1999) calculated ranges between 300 acorns per m2 to 25/m2. In dehesas, individual trees

produce an average of 12-18 kg of acorns per tree, but some may produce as much as 600

kg in a single year (Parsons 1962).

Quercus suber, the cork oak, prefers siliceous soils and 600-800m rainfall. It prefers

the lower elevations of southern Spain and Portugal but it is found at elevations up to

800m in northern Portugal. Cork oaks prefer mean annual temperatures of ~15° C but

will tolerate temperatures as low as -5° C. Acorns mature in autumn and drop between

October and March. Yields are slightly lower for cork oaks compared to the Holm oak.

The deciduous oaks are generally restricted to northern Portugal and upper elevations

in the central part of the country. Quercus faginea prefers the 600-1200 m elevation zone

but is still common in the lowlands. It will tolerate hot summers and temperatures as low

as -12° C in winter. Its acorns mature in September. Q. pyrenaica is mainly found between

400 and 1600 m asl, but can grow up to 2100 m asl. It prefers siliceous soils and annual

rainfall above 600 m. Its temperature range is -7° C in winter and 22° C in summer. Its

acorns mature in October and November. Q. robur is found between 0 and 1000m asl, but

as high as 3000m in northern Portugal and Spain. It requires at least 600 m annual rainfall

and is limited by winter lows of -15° C and summer highs of 25° C. Acorns mature in

September and fall in October.


Table : Macronutrient values for oak acorns from the Mediterranean and California
Species Fats Carbohydrates Proteins kcal/g
Eur ope
Quercus ilex     
Q suber    
Q robur  
Ca lif orni a
Q lobata      
Q garryana     
Q douglassi      
Q chrysolepis 
Q agrifola    
Q kelloggii     

Data from Basgall (
); FEDNA (Spain) ) Tabela da Composição dos Alimentos Portugueses
(Gonçalves Ferreira & da Silva Graça  )

Table 4.6 shows the nutritional characteristics of Q. ilex and Q. suber acorns compared

to the California oaks. In contrast to pine nuts, the acorns are poor in protein and fat, but

much higher in carbohydrate. Fat and protein content are comparable to the California

species but both are considerably lower on average in percent carbohydrate.

Recently, Harrison (1992, 1996) has brought attention to the highly productive and

“sweet” acorn of the holm oak. This species is widespread in Iberia due to the dehesa

(montado in Portugal) land-use system in which pigs are fattened on acorns prior to

slaughter. The two primary oak species in the dehesa/ montado are Q. ilex and Q. suber. For

Q. ilex, 9 kg of acorns translates into 1 kg of pork (Joffre et al. 1999). In some areas, pigs

and cattle will eat the acorns from Q. faginea, the Portuguese oak, however its acorn is

bitter compared to that of Q. ilex and Q. suber. The Pyrenean oak, Q. pyrenaica, is also

incorporated into the montados north of the Tejo. Pollen studies in southwest Spain suggest

the earliest physical record of such managed woodlands dates to about 6,000 bp (Stevenson

and Harrison 1992). While the dehesa/montado system is part of a pastoralist economy, it

does not necessarily require domestic plants or animals. The system enhances productivity

by encouraging oak groves as well as several other arboreal species in order to increase

browse for pigs, goats and cattle. In addition, these human-created ecosystems are home

to a wide diversity of wild animals including red deer, wild boar, rabbit, Iberian lynx and

birds (Fernandes de Abreu et al. 1993).

Hunter-gatherers incorporating simple forest management techniques such as

pruning, burning or possibly intentional planting could have created improved foraging

areas for wild boar, deer, chamois and even wild aurochs. Spring pruning in the dehesa/

montado is the primary method for increasing acorn yields per tree however this would be

difficult if not impossible to detect archaeologically. There is evidence of prehistoric fire

management of European woodlands by people during the Mesolithic (Mellars 1976; Mason

2000). Much of this burning has been perceived as a means of encouraging new growth

for browse to support deer and other ungulates. However, as Mason (2000) points out,

burning can encourage the proliferation of desirable forest species for human subsistence.

In this case, fire may have been used as a tool to manage oaks or other fruit/nut-bearing

vegetation. Fire may permit more light to reach the crown thus increasing acorn yield for

individual trees (Mason 2000). Comparisons between Holm oaks in managed stands and

natural forests showed that unmanaged trees are generally shorter, found closer together

and have smaller canopies (Pulido et al. 2001).

Fire also benefits the cork oak which is used today to reforest fire-prone areas because

of their thick protective bark (Carrión et al. 2000). It also produces what is considered a

“sweet” acorn, meaning that they are low in tannin. These sweet acorns of southern Iberia

require minimal roasting time to make them palatable. It should be noted that the amount

of tannin does not correlate well with perceptions of edibility. Some aboriginal groups in

California preferred the most bitter acorns, possibly due to their higher fat content (Basgall

1987; Mason 1994, 1995).

Lewthwaite (1982) has raised the possibility that humans played a deliberate role in

selecting sweet acorn varieties. This could have important consequences for lowering

processing costs to remove tannins prior to human consumption. On the other hand,

humans may have intentionally selected sweeter acorn varieties after observing that pigs

prefer the sweeter ones (Parsons 1962). Oak stands could have been managed to provide

carbohydrate and fat for human diets or to create animal fat and protein, ‘porridge or

pannage’ to borrow from Grigson (1982).

The emphasis here on pine nuts and acorns serves to illustrate the productive yield of

plants and their nutritional value regardless of processing costs. Other productive and

nutritionally valuable tree nuts may have been available such as chestnuts and hazelnuts.

Table 4.4 also shows the nutritional values for each of these along with pine nuts and


4.3 Late Pleistocene/ Early Holocene Portugal

The examples noted earlier provide evidence that plants were available and utilized

by prehistoric people throughout the Late Pleistocene and Early Holocene in Iberia. The

question arises with the timing of their availability and the antiquity of significant plant

exploitation in central Portugal. There is a nearly complete absence of macrobotanical

evidence of plant use during this period. Instead, inferences must be made based on

paleoenvironmental studies, including pollen and charcoal, interpretations of stone tool

technology, microscopic use-wear and recovery of starch grains. The last two are only

beginning and no definitive results can be presented here. Thus, the discussion will focus

on the potential plant availability from paleoecological studies, interpretations of stone

tool technology and expectations derived from experimental and ethnographic studies of

economic plant utility in other regions.

Climate change and paleoenvironment

Paleoenvironmental data for Tardiglacial and early Postglacial Portugal are extremely

limited. Information comes from geomorphology, deep-sea cores, pollen, archaeofaunal

and charcoal studies. Some of these data have been integrated into regional syntheses of

the Portuguese Upper Paleolithic by Bicho (1993, 1994) and Zilhão (n.d., 1997, 1998). The

standard interpretation was based on CLIMAP (1976), pollen diagrams from Lagoa

Comprida in the Serra da Estrela and nearby ponds in northwest Spain (Janssen and

Woldringh 1981; Turner and Hannon 1988; Mateus 1993, van den Brink and van Leewarden

1985, 1997), charcoal analysis from Cabeço de Porto Marinho (Figueiral 1993),

sedimentological and microfaunal evidence from Caldeirão Cave (Zilhão 1992; Póvoas et

al. 1989), and geologic context of open-air sites (Marks et al. 1994). New data published

since these syntheses are considered here.

The general circulation model of CLIMAP (1976) showed the Polar Front as low as

northern Portugal (42˚ N) during the Last Glacial Maximum (LGM), remaining there until

approximately 12,000 bp (Ruddiman and McIntyre, 1981; COHMAP 1988). The occurrence

of cold temperatures below 42˚ was also attributed by Bard et al. (1987) to the southward

Table a: Climatic chronology

Pollen phase Radiocarbon dates Oxygen Isotope
(uncalibrated) Stage
  bp Last Glacial
Lascaux Angles   bp
Termination A
Oldest Dryas (I)   bp Heinrich 
Bølling    bp

Older Dryas (II)    bp

Allerød    bp

Younger Dryas (III)    bp


Atlantic  bp

movement of the Polar Front. However, summer sea surface temperatures (SST) were

only a few degrees lower than present between 18-14,500 bp off southern Portugal

(Duplessy et al. 1992). The discrepancy between these studies is therefore explained by a

steep thermal gradient caused by the southward position of the Polar Front and the

movement of warm tropical waters northward. On the other hand, Fatela et al. (1994)

used benthic foraminifera data to argue that this cooling was due to the influx of cold

glacial meltwater into the North Atlantic during Termination IA of Oxygen Isotope Stage

2 deglaciation (Table 4.6a). Furthermore, they argue that the Polar Front never penetrated

south of 42°N during the LGM or deglaciation (Fatela et al., 1994; Abrantes et al., 1998).

Zahn et al. (1997) estimated SST for the LGM (20-17k bp) of 18-23˚C for winter and 22-26˚C

for summer due to oceanic circulation patterns that brought warm waters to the Portuguese

margin. Termination 1A is also marked by the occurrence of a Heinrich event. These

correspond to periods of iceberg flow from the Laurentide and Fenno-Scandinavian ice

sheets into the North Atlantic between 40°N and 60°N where they melt and the dust that

was locked in the ice accumulates on the ocean floor (Abrantes et al. 1998; Baas et al. 1998;

Grousset et al. 2000; Chapman et al. 2000). The occurrence ofsuch “ice-rafted detritus”

(IRD) in cores off the coast of Portugal indicate the southernmost drift of icebergs during

Heinrich events (Lebreiro et al. 1996). Not surprisingly then, the deglaciation is marked

by several fluctuations in temperature and salinity.

Duplessy et al. (1986) argued that continental deglaciation of northern Europe took

place in two phases, the first (Termination 1A) occurred 16,000-13,000 bp and the second

10,000-8,000 bp (Termination 1B), with no apparent melting 13,000-10,000 bp. Other studies

placed the initial deglaciation around 14,000 (Duplessy et al. 1986). Later studies of oxygen

isotopes and planktonic foraminifera from deep sea cores off southern Portugal by Bard et

al. (1987) show February sea surface temperatures (SST) as low as 4°C (15°C today) between

14,500-12,500 bp. Duplessy et al. (1996) show 11°C SST (21°C today) for summer. Both of

these studies would appear to agree that conditions were much colder during the

Termination 1A/H1/Dryas I in Portugal. The cause of the lowered SST and overall salinity

has been attributed to meltwater influx from the continental ice sheets. After Dryas I,

summer SST was as high as today during the Bølling/Allerød. In core SU 81-18 the Younger

Dryas is characterized by a rapid drop in SST and salinity (Duplessy et al. 1992).

Geochemical analyses of sediment in Caldeirão Cave Layer Eb suggest strong humid

(warm?) winds blowing across Estremadura some time after 16,000 bp and before 10,000

bp (Cruz, 1992). However, the lack of stratigraphic and chronological resolution makes it

impossible to know when this occurred. It may correspond to the Lascaux or Bølling/

Allerød interstadial. Zilhão’s (1997) interpretation of the Caldeirão stratigraphy suggests

the latter.

During the Last Glacial Maximum, sea levels were approximately 140m below their

current level (Dias 1985; Dias et al. 1997; Rodrigues et al. 1991). In some places along the

Estremaduran coast , the LGM shore was approximately 40 km east of its current position.

By about 16,000 bp, sea level rose to about -100m but large areas of land were still exposed,

up to 30 km in much of Estremadura. According to Dias et al. (2000) sea levels remained

steady until about 13,000 bp. Between 13,000 and 12,000 bp, sea levels rose to about 40m

below their current level, effectively shrinking the available land surface of Estremadura

by about 25% during the Tardiglacial (Figure 4.2). This corresponds to the beginning of

the Bølling/Allerød Late Glacial interstadial. During the Younger Dryas cold snap sea

level lowered again to 60m below present. Afterwards, between 10,000-8,000 bp sea level

reached 30m below present. This was followed by the Atlantic period transgression (7,500

bp) that created a large estuary in the lower Rio Tejo. By 3,500 bp sea level had stabilized

at its present level. While seemingly dramatic by geologic time, human groups would

probably not have faced crises in adaptation because the changes in sea level occurred

over hundreds of human generations (Waselkov, 1987). However, the lag between the

marine and terrestrial records of 180-350 radiocarbon years suggests more rapid shifts

(Boessenkool et al. 2001). Whatever the case, the submergence of about 40km of continental

shelf has, in all likelihood, drastically affected the archaeological record for the Solutrean

and Magdalenian in Portugal (Figure 4.2).

Information on vegetation in Portugal during the period 15,000-10,000 BP comes from

pollen studies in the Serra da Estrela and charcoal analyses from Cabeço de Porto Marinho

and Lapa do Suão (Van der Knapp & Janssen, 1991; Figueral, 1993; Haws and Valente,

2001). Pollen evidence from Charco da Candeeira, a pond located at 1400 m asl in the

Serra da Estrela, shows a rapid succession of steppe grasses and composites to herb and

Ericaceae near the pond in the early Bølling (Figure 4.3). The occurrence of pine, birch

and oak forest pollen suggests an expansion of forest refugia at lower elevations during

this interstadial, dated 12,600 bp (Van der Knaap and van Leeuwen 1997). Prior to this,

pollen was almost entirely absent possibly due to the outwash of montane glaciers.

Unfortunately, no other pollen cores exist in Portugal earlier than 12,600 bp. In fact there

El Juyo
La Riera
Altamira Rascaño


Atlantic Ocean

Cova Fosca
Buraca Grande
Lapa do Picareiro Caldeirão Cova dels Blaus
Bocas Cueva de la Cocina
Lapa do Suão
Volcán de Faro
Les Mallaetes
Parpalló Santa Maira
Tossal de la Roca
Cova de les Cendres

Vale Boi

Nerja Mediterranean Sea


Figure  : Map of Iberia showing the coastline (heavy line) during the Late Magdalenian
The faint line is the present coastline Note the amount of lost land in central Portugal
and Mediterranean Spain compared to Cantabria

Laguna Lucenza
Brañas de Lamela Quintanar de la Sierra
Mougás Sanabria Marsh Banyoles

N Lagoa de Marinho

Charco da Candeeira
Lagoa Comprida
Chafariz do Rei
Atlantic Covão do Boieiro


Fernão Ferro

El Asperillo
San Rafael Mediterranean Sea


Figure  : Map of Late Pleistocene pollen core locations mentioned in the text

are no low elevation pollen cores for the Late Pleistocene. After the interstadial forest

expansion, the Charco da Candeeira core shows a climatic downturn corresponding to

the Older Dryas (Dryas II) phase in northern Europe. This was followed by an amelioration

correlated with the Allerød interstadial. In the nearby Covão do Boieiro deciduous and

evergreen oaks increase approximately 10,000 bp (van der Knapp and van Leeuwen 1997).

An additional 60cm of undated Late Glacial deposits shows a predominance of Pinus

sylvestris/pinaster type pollen with a reduced but constant occurrence of oak and minor

contributions of birch, alder, sycamore, ash, hazel, olive and yew. The authors place these

deposits in the Allerød and Younger Dryas phases. Thus it appears that trees were

widespread in the mountains by 11,500 bp. A number of shrubs are also evident including

Calluna, Erica, Genista, and Juniperus. Their presence, along with grasses, suggests an open

woodland vegetation that reached a peak in density prior to the Younger Dryas. No

radiocarbon dates have been made, but the Chafariz do Rei sequence (1770m) shows Pinus

and Artemisia at the base possibly corresponding to the Bølling (van der Knapp and van

Leeuwen 1997). Oak and birch appear during the late Bølling/Dryas II/Allerød sequence.

At a slightly lower elevation (1150m), the Lagoa de Marinho sequence dates to the Allerød

phase. The arboreal pollen is dominated by deciduous oak with lesser amounts of evergreen

oak, hazel, birch, chestnut, holly, ash, alder, poplar, willow and myrtle (Ramil Rego et al.


From a regional perspective, additional pollen studies have been made in adjacent

areas in Spain, namely Galicia and Andalucía. Galicia is important because it (and northern

Portugal) marks a biogeographical boundary between the Euro-Siberian and Mediterranean


zones (Muñoz Sobrino et al., 1997). Cantabria is considered entirely within the Euro-Siberian

belt. Andalucía is fully Mediterranean today. The only pollen sample in western Iberia

dated to the Late Pleniglacial comes from Laguna Lucenza (1375 m asl) in Galicia (Muñoz

Sobrino et al. 2001). Cold loving and sun-loving plants dominate the period before 17,300

cal yr BP. Trees present include pine, birch and deciduous oak. Dryas I (here dated

17,300-15,300 cal yr BP) is marked by an increase in Artemisia indicating cool, dry steppe

conditions. However, tree pollen in low percentages includes pine, juniper, birch, hazel

and deciduous oak. Between 15,300 and 13,300 cal yr BP arboreal pollen expands to as

high as 80%. Added to the previous types are evergreen oak, chestnut, alder and ash. On

the coast at Mougás, tree pollen is low with dominance by humid-adapted heathers and

grasses (Ramil Rego et al. 1998). Additional pollen studies in the mountains of northwest

Iberia show a Bølling/Allerød vegetation composed of 40–60% trees, mainly pine and

birch with some deciduous oak, hazel and chestnut (Muñoz Sobrino et al. 1997). Grasses

and herbs made up the rest. The sequence from Brañas de Lamela (1280 m asl), a more

south-facing, sheltered basin, contained the Mediterranean-type evergreen oak (Quercus

ilex). It would appear therefore, that climate during this period was very similar to today.

In southern Spain, pollen cores dating to the Late Pleistocene have been published

from El Asperillo (0-30m) near Huelva, Padul (785 m asl) near Granada, San Rafael (near

Almería (Pantaléon-Cano et al. 2003) and Navarrés (225 m asl) near Valencia (Stevenson

1984; Pons and Reille 1988; Carrión and van Geel 1999). The El Asperillo sample is from a

peat band dated to 13,000 bp. It shows a slightly cooler and wetter climate with a downward

elevational shift in forest cover of about 100 m compared to today. Several studies have

been conducted at Padul, the most recent supported by 21 radiocarbon dates (Pons and

Reille 1988). The LGM appears to be a time of widespread arid steppe vegetation with

grasses and some pine trees. Oaks and Ericaceae are present but in very low percentages,

indicating a nearby refugium of Mediterranean species. About 15,000 bp, pine pollen

decreases sharply as steppe species increase, suggesting a possible climatic amelioration.

Pons and Reille (1988) conclude that climatic amelioration did not take place until after

13,000 bp, when evergreen oak (Quercus ilex) pollen increases. The core from the Canal de

Navarrés (Valencia) shows high aridity at 18,000 bp with a gradual replacement of Artemisia

by pine sometime afterwards. The Dryas II is recorded at 12,000 bp by a return of Artemisia.

Its occurrence in several peaks suggests rapidly fluctuating climatic conditions during the

Tardiglacial (Carrión and Van Geel 1999). At San Rafael, the Upper Pleniglacial vegetation

is marked by a high arboreal pollen percentage (~75%) (Pantaléon-Cano et al. 2003). The

most abundant tree pollen comes from evergreen and deciduous oaks wild olive, with

mastic also evident. Grasses are fairly well represented indicating perhaps an open


Recently, marine pollen records from deep sea cores have been published

(Hooghiemstra 1988; Hooghiemstra et al. 1992; Parra 1994- cited in Carrión et al. 2000).

Pollen diagrams from cores SU 8103 off SE Spain, SU 8113 off SW Spain and 8057B off SW

Portugal show interesting correlations to the terrestrial cores. In all three, pine dominates

the arboreal spectrum but other deciduous and evergreen trees are present from the LGM

onwards. After 14,900 in core 8057B, Pinus, Artemisia and Chenopodiaceae decrease sharply

at the expense of evergreen and deciduous oak (Hooghiemstra et al. 1992). In the cores off

Spain, there is a more gradual shift from the Artemisia and Chenopodiaceae to the evergreen

oaks around 12,000 bp (Carrión 2000). Sánchez Goñi et al. (1999, 2000, 2002), Boessenkool

et al. (2001) and Roucoux et al. (2001) have correlated the terrestrial pollen records with the

marine record from the Last Interglacial to present. Boessenkool et al. (2001) show the

spread of deciduous oak slightly later, around 13,000 bp, than the record from core 8057B

nearby. Turon et al. (2003) show the same pattern for core SU 81-18 to the west of 8057B In

core MD95-2039, a site 180 km offshore to the west of the mouths of the Douro and Mondego

rivers, arboreal pollen reaches higher than present levels after Heinrich I (Roucoux et al.


The general picture for the Last Glacial Maximum climate in Iberia is of aridity with

slightly cooler temperatures followed by instability between 17-14,500 bp. After a cold

event triggered by meltwater and iceberg rafting around 14,500 bp, temperatures and

precipitation increased by13,000 bp, allowing forests to return to the uplands. The lack of

low elevation pollen records for much of Iberia precludes a definitive paleoenvironmental

reconstruction. In all cases, except El Asperillo, Mougás and San Rafael, pollen cores

come from lakes, ponds or peat deposits in mountain zones. Most have no record of the

initial deglaciation or at best imply a cool arid steppe vegetation prior to the last glacial-

interglacial transition. In spite of this problem, warm adapted Mediterranean trees are

evident in these cores albeit in very low frequency. This likely reflects a regional pollen

input and that low elevation zones must have been refugia for these plants (Carrión 2002).

The enhanced aridity evident from pollen cores during the Heinrich events was likely

caused by shifts in atmospheric circulation patterns. The present-day Mediterranean


: Pollen sites in Iberia with cores extending to the Late Pleistocene

Location setting elevation basal age

NW Portugal
Charco da Candieira mountain pond    /
Lagoa Comprida mountain lake    /

Lagoa Clareza mountain lake    /
Covão do Boieiro mountain pond 
 Late Glacial
Lagoa de Marinho mountain lake    /

SW Portugal
Fernão Ferro coastal lagoon   

NW Spain
Laguna Lucenza mountain lake  
Queixa Sierra mountain   
Sanabria Marsh mountain marsh     /
Laguna de la Roya mountain lake  
Mougás coastal lagoon   / 
Brañas de Lamela mountain bog   Late Glacial
Lago de Ajo mountain 

Southern Spain
Navarrés flat valley   / 
Siles mountain lake   
Padul mountain
El Asperillo coastal bog    

climate regime is characterized by hot dry summers and mild wet winters. During the

LGM and Termination 1A/Heinrich 1/Dryas I the winter precipitation decreased because

of increased stability in high-pressure due to the lack of southward movement of polar

highs (Combourieu Nebout et al. 2002). Conversely, during millennial-scale oscillations

of interstadial conditions known as Dansgaard-Oeschger events higher winter precipitation

and summer temperatures permitted forest expansion (Combourieu Nebout et al. 2002;

Sánchez Goñi et al. 2000, 2002). The slow rate of expansion of oak forests and the long lag

between the marine pollen and SST records suggest that warming began much earlier

than 13,000 bp (Boessenkool et al. 2001; Chapman et al. 2000).

Charcoal analyses at the Upper Paleolithic site of Cabeço de Porto Marinho provide

the only low elevation Tardiglacial paleobotanical data for Portugal. Figueral (1993)

identified components of a mixed Atlantic/Mediterranean community of pine (Pinus

pinaster/ pinea), evergreen and deciduous oak, birch (Betula pubescens), olive (Olea europaea

var. sylvestris), ash (Fraxinus angustifolia), wild strawberry (Arbutus enedo), tree heath (Erica

arborea) and Leguminosae in CPM levels dated 11,200 bp, corresponding to the Allerød.

Charcoal analysis of the Magdalenian levels of Lapa do Suão shows a similar composition.

Species identified include deciduous oak, pine, wild fruit (members of the Rosaceae family)

and olive. Radiocarbon inversions preclude a high resolution chronology of plant

succession. Lapa do Picareiro has abundant charcoal in levels dated between 8,310-12,300

bp but it has not been fully analyzed. Preliminary analysis points to a similar vegetation

comprised of deciduous oak, pine, wild olive, willow, ash and Leguminosae typical of

Mediterranean garrigues. The majority of oak in the Late Magdalenian levels is a deciduous

oak, possibly Q. faginea. The dominance of this species in the Magdalenian levels indicates

warm, humid conditions as its highest densities in Iberia today occur along the

Estremaduran coast down to southern Portugal.

In Spain, charcoal analyses have been done on numerous sites in Cantabria, the

Pyrenees and the Mediterranean region. Late Glacial assemblages from Cova de les

Cendres, Cova de Santa Maira, Ratlla del Bubo, Tossal de la Roca, Cova Bolumini and

Cueva de Nerja in the Mediterranean region show a much greater abundance of trees

during the Dryas I phase 15,000-12,500 bp than pollen diagrams. This is likely due to

elevational differences between the montane ponds and lakes where the pollen cores were

taken and the archaeological sites. At Ratlla del Bubo, dated to the Solutreo-Gravettian,

juniper dominates but several other trees are present in low frequency. These include

turpentine tree and mastic (Pistacia terebinthus and lentiscus), wild olive (Olea europaea var.

sylvestris), fig (Ficus) and ash (Fraxinus oxyphylla) (Badal and Carrión 2001). Charcoal from

the preceding Solutrean levels at Cendres was predominately black pine (Pinus nigra),

with woody Leguminosae, juniper and small percentages of the evergreen oaks, Quercus

ilex and Q. coccifera (Villaverde et al. 1999). Herbs and shrubs were also identified. During

the initial warming, pine decreased as juniper, Leguminosae and evergreen oak increased.

Also identified during this period are Prunus sp., deciduous oak, heather (Erica multiflora)

and Ephedra. The Dryas I cold phase is marked by a sharp increase in juniper at the expense

of pine and Leguminosae. The evergreen oaks decrease but are still present. In the Upper

Magdalenian levels corresponding to the Bølling/Allerød interstadials pine comprises

over 50% of the assemblage with juniper, Leguminosae and evergreen oak in low

percentages. At the end of the occupation deciduous oak, poplar and Pinus halepensis

appear. Villaverde et al. (1999) note that the environment surrounding the shelter was

probably as dry as today but slightly cooler due to the co-occurrence of black pine and

evergreen oak. At Tossal de la Roca the early occupation level dated 15,360 bp had charcoal

from pine (Pinus nigra), with low percentages of juniper and deciduous oak (Quercus

faginea), and box (Buxus sempervirens) (Cacho et al. 1995). This suggests a similar vegetation

to Cendres for the same period. In the Final Magdalenian level dated after 12,500 bp pine

still dominates but deciduous oak increases from 1% to 18%. Montpellier maple (Acer

monspessulanum) and rockcherry (Prunus mahaleb) appear in low frequencies. In the

Microlaminar Epipaleolithic level (10,500) pine drops to a low percentage while oak and

juniper dominate. In the Exterior section, the dated to the Geometric Epipaleolithic, here

dated 9,000-7,000 bp, there is a higher diversity of trees represented in the charcoal

assemblage. Evergreen and deciduous oaks comprise about 80% of the assemblage

followed by Prunus mahaleb, Sorbus sp.(another wild fruit tree), maple and pine (Cacho et

al. 1995). The Santa Maira charcoal assemblage shows a steady decline in juniper from

about 60% in the Upper Magdalenian levels to less than 10% by the Geometric Epipaleolithic

(Badal and Carrión 2001). During this time evergreen and deciduous oaks comprise about

50% followed by low percentages of Prunus, maple and poplar. Other Mediterranean

types include ivy (Hedera helix), Cistacae, Erica sp., Fraxinus sp., Buxus sempervirens and

Pinus pinea. The vegetation represented is similar to that identified at CPM in Portugal

during the same period with the exception of juniper. This may simply be due to elevational

differences between the two sites. At Bolumini, the brief Late Magdalenian levels show

the oak expansion during the Bølling/Allerød interstadial.

Microfaunal analyses are also useful in accurate paleoenvironmental reconstruction.

Together with pollen and charcoal studies, these data can help balance the biases in regional

pollen inputs and human selection of wood fuel. In Portugal, Póvoas et al. (1992) argue

for a shift to a Mediterranean climate during the formation of Level Eb at Caldeirão, dated

14,450-10,700 bp. This interpretation is based on the increase in percentages of the

Mediterranean pine vole Terricola duodecimcostatus and decrease in the ratio of the voles,

Microtus arvalis to M. agrestis. Unfortunately the rodents are mixed in with ceramics and

sheep bones dated directly to 6,200 bp making it impossible to recognize climatic

fluctuations seen in the deep-sea record. At Picareiro, during the Upper Paleolithic

sequence, Markova (n.d.) notes high percentages of Terricola duodecimcostatus together

with Apodemus sylvaticus (field mouse) and Eliomys quercinus (garden dormouse) indicative

of Mediterranean forested environments and rocky habitats in Level J. This trio dominates

the entire sequence with subtle changes observed in Level I where the snow vole, Chionomys

nivalis, appears and Level F where Microtus agrestis occurs. Level I could date to either the

LGM, Upper Pleniglacial or Dryas I which would have been times of cooler and relatively

dry conditions. This is supported by the deposition of eboulis sec in the cave. Level F dates

almost entirely to the Dryas II climate phase which was likely more moderate (wetter)

than today as indicated by the presence of Microtus agrestis (Markova, n.d.). Indeed, Dryas

II barely registers in the deep sea core (SU81-18) off the coast of southern Portugal and is

absent in many regional pollen cores (Bard et al. 1987; Peñalba et al. 1997; van der Knapp

and van Leeuwen 1997; Carrión and Van Geel 1999). The presence of the water vole,

Arvicola terrestris, in E Lower is not surprising given the radiocarbon overlap between this

level and Level F. In the Magdalenian levels of Lapa do Suão, dated to roughly 15,000-

10,000 bp, Haws and Valente (2001) identified Eliomys quercinus, Arvicola terrestris and

Microtus agrestis. The composition matches that of Caldeirão Eb and Picareiro E and F.

Pleistocene paleoenvironmental reconstructions also rely on macrofaunal remains

from archaeological sites (Bicho, 1993, 1994; Cardoso, 1993a &b; Zilhão, 1995). At Caldeirão,

the disappearance of ibex (Capra pyrenaica) and chamois (Rupricapra rupicapra) and

subsequent replacement by red deer, roe deer, wild boar and beaver has been argued to

indicate a change from cold, dry steppe to warm, humid forest conditions. Explicit in this

argument is the idea that the former animals are cold-adapted because of their presence in

alpine areas of Iberia today. However, researchers in Mediterranean Spain use chamois

as an indicator of warm, humid forest conditions in the early Holocene. Although chamois

today is usually found in higher elevations across Europe, Tosi et al. (1987) show it living

at 300m elevation in Italy. These facts support the idea that chamois have a much restricted

range than in the past. Indeed, Miracle & Sturdy (1991) presented evidence from the

Balkans that habitat alteration and human hunting pressures in the recent past have forced

chamois out of low-elevation karst zones. Therefore, the disappearance of chamois from

the Caldeirão assemblage likely represents a change in human selection of this prey, not

simply a change in climate. Its occurrence in Allerød levels at Picareiro provides further

support (Haws, 1998).

Although data are scanty it is likely that the Younger Dryas impacted central Portugal.

Bard et al. (1987) and Duplessy et al. (1996) show cold waters of the coast of southern

Portugal during the Younger Dryas. Magnetic susceptibility curves for Picareiro show a

sudden drop between 11,700 and 10,070 bp likely corresponding to the Younger Dryas

(Ellwood et al. 2001). An abrupt decrease in oak pollen from marine cores suggests a

rapid decline in temperatures on land. Pollen spectra from northern and southern Spain,

however, show a return of steppe communities indicative of dry but not very cold

conditions (Pons & Reille, 1988; Peñalba et al. 1997; Carrión and Van Geel 1999). Both

evergreen and deciduous oaks remain present in Padul and Quintanar de la Sierra (1470

m asl) but in lesser frequency. In Catalunya, the Lake Banyoles record shows a drop in

Pinus but other species remained constant (Pérez-Obiol & Julià, 1994). The molluscan

evidence from Pedra do Patacho dated 10,400 bp have been argued to indicate cold waters

off the southwestern Portuguese coast due to the presence of Littorina littorea. However,

these limpets are found today on the Algarve coast casting doubt on their use as

paleoclimatic indicators.

In the early Holocene, by 9,500 bp, a mix of Mediterranean and Atlantic species

returned to Estremadura. Pollen diagrams from the Lagoa Comprida (1400 asl) show

vegetation at 9,200 bp dominated by pine with Artemesia and Chenopods indicating an

open forest (Jansen and Woldringh, 1981). Between 9,200-9,000 bp, there was a rapid

transition from the open pine forest to a more closed one dominated by oak. Shortly

before 8,300, birch pollen rises along with grasses. Between 8,300-2,600 bp, the pollen

record shows several oscillations of oak and birch with small percentages of pine. Ash,

willow, hazel, and alder are also present. On the coast, the earliest pollen record is dated

9,500 bp just south of Lisbon in the Lagoa do Golfo (Mateus and Queiroz 1997). Since the

base of the core was dominated by pine, Mateus and Queiroz (1993) argue for extensive

Maritime pine (Pinus pinaster) stands on the coastal dunes. They extrapolate this hypothesis

further back to the Late Glacial and suggest the entire coastal strip from Sines in Alentejo

to Porto was covered by Maritime pine. Obviously, this requires data from offshore since

the Late Glacial coast lay 10-40 km west of its present location.

Many researchers have attempted to correlate the climate changes documented in

deep sea and ice cores with terrestrial records and subsequently argue for similarities

between the changes in northern and southwestern Europe (de Beaulieu et al. 1994; Lowe

and NASP Members 1995; Peñalba et al. 1997; Zilhão 1997;. Roucoux et al. 2001; Ellwood et

al. 2001). Indeed, there are agreements and disagreements between the different records.

However, the evidence from Portugal shows a much more moderate climate (similar to

today) during the last glacial-interglacial transition than in Northern Europe or even

northern Spain. While the Polar Front may have been situated off the coast of Galicia

during the Last Glacial Maximum and montane glaciers grew in the mountains of north

and central Portugal, climate was never severe enough to permit colonization by cold,

steppe-adapted mammoth and reindeer as in Cantabria. The plant and animal communities

in Portugal were flexible enough to adapt with some altitudinal and latitudinal shifts

resulting in localized changes in community composition, mainly the proportions of species

represented. Thus, non-arboreal shrubs and sedges expanded their range in the mountains,

but trees were still present. Mediterranean types shifted downward in elevation and

possibly southward, while birch, pine and deciduous oak prevailed in montane valleys.

Horse and wild aurochs likely had increased area of grassy plains to grow their populations.

Ibex and chamois may have been more numerous to the detriment of red deer and wild

boar populations. By 17,800 bp sea surface temperatures rebounded to those of the present

day. Unfortunately, there is no solidly-dated terrestrial record in Portugal for the period

between 18,000 and 12,500 years bp. One can only assume conditions on land also

improved. The first indications from charcoal analysis show this to be true at least by

14,000 bp. The composite magnetic susceptibility reference section for southern Europe

by Ellwood et al. (2001) shows the Lascaux interstadial followed by two warm oscillations

prior to the Allerød. It is likely therefore that the warm ocean temperatures dated 17,800

bp correspond to the Lascaux interstadial. The second oscillation reflects the onset of the

Bølling period. The Younger Dryas cold snap seems to have resulted in cooler and more

arid conditions across much of Iberia.

Reconstructing plant availability for Late Pleistocene Portugal is also made difficult

by the millennia of landscape alteration by people since the Neolithic. Recent planting of

Maritime pine (Pinus pinaster) forest since the 12th century has led to an artificial abundance

of this tree species in central Portugal (Vieira et al. 2000). This species has great economic

value for its resin and wood, which was used to build ships. Pinus pinea was also valued

in ship building and it was more widespread in the historic past. During the Arab

occupation, dense forests of P. pinea were cleared in order to export wood to North Africa

and the Middle East (Vieira et al. 2000). The planting of Stone pine plantations in the

south has also led to the common idea that they are better adapted to warmer, sunnier

climates while the former prefer slightly more temperate conditions. In protected natural

places in Estremadura like the Tapada de Mafra (the former royal hunting grounds) both

pines are present. In general P. pinea outcompetes P. pinaster on the sunnier, south-facing

slopes of hills while on the northern face the opposite is true. Identifying their occurrence

in prehistoric archaeological sites can be problematic since ecologists consider them both

part of the meso-Mediterranean (Barberó et al. 1998). Charcoal studies are perhaps the

most common method used. Figueiral (1995), Schweingruber (1990), Jacquiot (1955) each

give criteria for distinguishing the two types based on wood anatomy. Mateus and Queiroz

(1997) identified Pinus pinaster charcoal from a single hearth at Ponte da Vigia and pollen

cores from Fernão Ferro, leading them to extrapolate a Maritime pine forest on the Late

Glacial coast of Estremadura. Given small sample size, single species dominance, and the

fact that both date to the early Holocene one has to wonder about the accuracy of this

reconstruction. Zilhão’s (1997) hypothesis that the Late Glacial coast was covered by sand

dunes is based on four lines of evidence: 1) the Solutrean site of Vale Almoinha, which

today is located near the coast in eolian deposits; 2) sedimentary analyses of submerged

landforms by Daveau (1980) that suggest desert-like condistions on the LGM coast; 3) the

pollen/ charcoal record from Fernão Ferro; and 4) the geochemical study of Caldeirão Eb

showing elevated Na, argued by Zilhão to indicate the persistence of cold, salty winds

blowing inland. Vale Almoinha is probably 40 km inland from the LGM coast and therefore

should not be considered a “coastal” site. Therefore conclusions about the origin of the

eolian deposits need to be reconsidered. The pollen record from Fernão Ferro is reported

to have a high percentage Pinus pollen which is assumed to be P. pinaster. Caldeirão Eb is

badly mixed and the two radiocarbon dates are on bulk bone samples. There is no reason

to conclude from them that the deposit accumulated gradually over 5,000 years, nor that

elevated Na in the deposit is due to salty winds blowing inland from the ocean, especially

given that the cave is located in a sheltered valley 60 km inland. Furthermore, Turner and

Hannon (1988) suggest that deciduous forest refugia during periglacial times were likely

the maritime coasts of Spain and Portugal.

As evident in the pollen and charcoal data, southern Spain and central Portugal

witnessed a much earlier expansion of warm, temperate and Mediterranean forests after

the LGM than in northern Spain and Catalunya. The Dryas I phase saw a regression of

forest species. In southern Spain and central Portugal the reappearance of mixed temperate

and Mediterranean vegetation took place in the Bølling/Allerød phase whereas in northern

Iberia it occurred in the Early Holocene. Rich plant communities were certainly available

to Magdalenian hunter-gatherers in central Portugal. It is necessary to consider the potential

utility of the available plants for Late Upper Paleolithic hunter-gatherers in central Portugal

and the factors that condition their deposition and preservation in archaeological sites.

4.4 Modeling economic and nutritional utility of plants

Researchers using energy-based foraging models to understand plant exploitation

among hunter-gatherers often implicitly ask the question, ’what are the costs relative to

the benefits of exploiting a given plant?’ With regard to many plants, especially tannin-

rich ones requiring high processing costs, the answer is usually that the costs often far

exceed the energetic benefits and that their inclusion in the diet must be due to some sort

of subsistence stress brought on by population-resource imbalance. However, an important

question may be asked from a wider nutritional perspective: ‘what are the costs of not

utilizing certain plant resources?’ Given the issues regarding human nutrition in chapter

2, all resources are not equal and probably should not be measured by a single variable,

energy, despite its appeal to simplicity (e.g., Jochim 1998). Non-energetic nutritional needs

can alter the perceived “value” of a food resource. Without relying on cumbersome linear

programming models that require extremely fine-grained data, plant (and animal)

resources can be evaluated with regard to nutritional quality for analytical purposes.

Most of the work on modeling prehistoric plant use has been done within a foraging

theory framework. The majority of these studies are centered in the Great Basin and

California, though models also have been developed for eastern North America (Simms

1987; Barlow and Metcalfe 1996; Gardner 1997; Gremillion 2002). Basgall (1987) considered

the role of acorns in prehistoric California subsistence economies. Simms (1987)

experimentally derived energetic return rates for wild plant resources utilized by prehistoric

Great Basin foragers. Note that Table 4.8 shows that plants and small game consistently

rank lower than large game. For Europe, Rowley-Conwy (1984) has also estimated return

rates for acorns and hazelnuts as well as other economic resources based on Perlman’s

(1980) figures for eastern North America (Table 4.9). However, neither Rowley-Conwy

nor Perlman considered handling times in their return rate estimation. Additional studies

focused on nutritional returns from wild plant collecting have been conducted in northern

Mexico and northwest Patagonia (Laferrière 1995; Ladio and Lozada 2000). These studies

show that energetic return rates are often not the primary factor determining plant food

choice by hunter-gatherers. Emphasis is given here to studies in western North America

and Europe that focus on acorns and pine nuts because these resources from different

Table  : Comparative return rates for plant and animal resources in the Great Basin
species common name/type kcal/hr

Anabrus simplex grasshopper  

Odocoileus hemionus mule deer 

Ovis canadensis bighorn sheep 

Antilocapra americana pronghorn antelope 
Lepus sp jackrabbit  

Thomomys sp gopher  

Sylvilagus sp rabbit   
Typha latifolia cattail (pollen) 
Spermophilus sp squirrel   
Citellus sp squirrel 
Anas sp duck 

Quercus gambelli acorn 
Descurainia pinnata tansymustard seed  

Pinus monophylla piñon seed 

Lewisia rediviva bitterroot 

Elymus salinas wild rye seed  

Atriplex nuttalli shadscale seed  
Atriplex confertifolia shadscale seed 
Scirpus sp bulrush seed  
Echinochloa crusgalli barnyard grass seed

Lepidium fremontii peppergrass seed 

Helianthus annus sunflower seed 

Poa sp bluegrass seed  
Elymus salinas wild rye seed 

Oryzopsis hymenoides ricegrass seed 
Phalaris arundinacea reed canary grass seed  
Muhlenbergia asperfolia scratchgrass seed   
Hordeum jubatum foxtail barley seed  

Carex sp sedge seed 
Typha latifolia cattail root   

Scirpus sp bulrush root  

Distichlis stricta saltgrass seed 

Allenrolfea occidentalis pickleweed seed 
Sitanion hystrix squirreltail grass seed 
Gila bocolor minnow

Data from Simms (


Table : Resource return rates suggested

by RowleyConwy ( )for Mesolithic Denmark
resource kcal/hr


terrestrial mammals 
sea mammals ?
waterfowl ?
fish ?
Return rates are based on collection rates only
Handling time was not considered

species of oaks and pines would have been present in the Late Pleistocene of central

Portugal. In fact, they are evident from Late Pleistocene and Early Holocene sites in

Mediterranean Spain.

For the Great Basin, Metcalfe and Barlow (1992) built a model for field processing

and transport of pine nuts from Pinus monophylla. This pine is widespread in the Great

Basin and was the most widely exploited variety in prehistoric and historic times (Rhode

and Madsen 1998). In the Metcalfe and Barlow model, field processing decisions are

conditioned by the travel time to the resource, its utility (edible: inedible portion) and the

time necessary to process it. Utility indices for pinyon pine nuts were then applied to a

central place foraging model to “explore relationships between the costs and benefits of

collecting, field processing and transporting loads …to base camps and the implications

for overall efficiency while foraging” (Barlow and Metcalfe 1996: 352). Bettinger et al.

(1997) also used a central place foraging model to explore acorn and mussel use in central

California. In each of these models, knowledge of distance and/or travel time to resource

location and the time needed to process are critical elements. The results of these studies

are used in this section to explore some of the factors that may have relevance to Late

Pleistocene and Early Holocene plant exploitation.

Unfortunately, paleoenvironmental reconstructions are not fine-grained enough to

know the locations of various resources in relation to an archaeological site. Often, site

types (i.e., residential vs task sites) are not easily determined. Given these problems the

central place foraging models cannot be fully utilized for most prehistoric cases. They do

however, provide some expectations concerning resource use that can be applied to Late

Pleistocene Portugal. The Metcalfe and Barlow (1992) model predicts when resources

should be processed in the field prior to transport. The feasibility of processing is dependent

on the type of ‘package’ the resource is found, or the inedible fraction that must be removed.

Removing this portion increases the utility of a resource and allows more to be collected

and transported back to camp. For plants, the greater the processing time required the

less likely a resource will be field processed. Time and group composition also place

important constraints on the decisions. If women and children are collecting, they are less

likely to camp overnight than men, thus placing a time constraint on collecting. Weight is

also an important variable because of thresholds in the amount that can be carried.

Reducing or eliminating bulky, inedible parts lowers the weight and increases the overall

utility of a resource.

In their experimental model for pine nut and pickleweed exploitation, Barlow and

Metcalfe (1996) found that foragers could lower costs and raise return rates if they moved

residence to the resource location. For pine nuts, the cones represent a significant, space-

consuming fraction and should be removed to maximize the amount of edible nuts

transported. The hulls require more processing and it is not economical to remove them

prior to transport unless extremely long distances are involved (>100 km). Unless stands

are within a very short distance pine nuts should be transported to base camps in their

hulls. This means that cones should be rarely deposited in residential camps, but pine nut

hulls may be deposited in large numbers. Since they do not require heating to be cracked

open, these may rarely be charred and therefore not likely to preserve except in rare cases

of exceptional preservation. Based on the central place model, if pine nuts were a significant

resource then residential camps would not necessarily be expected in areas near stands

because pine nuts can be transported fairly long distances before it becomes unprofitable.

In the Great Basin example, significant weight differences were found in the amounts

of processed vs unprocessed pine nuts that could be carried using basket containers whose

function as pine nut carriers is known ethnographically. Barlow and Metcalfe (1996) report

that these baskets could be filled with 6 kg of unprocessed cones vs 18 kg of processed

nuts. Of course, carrying such a large load over long distances would be physically

demanding. Ethnographic data suggest women would carry 3-15 kg (Metcalfe and Barlow

1992). Based on this alone it would seem doubtful that prehistoric foragers would have

traveled long distances to carry heavy loads. However, Barlow and Metcalfe (1996) show

that return rates are higher for larger load sizes regardless of processing time and distance

to patches. This pattern appears to hold true in other regions such as northwest Patagonia

where the Mapuche make overnight trips to the forest 50 km away to collect seeds of the

Araucaria araucana. Ladio and Lozada (2000) do not report individual load size, numbers

of carriers or whether cones were discarded prior to transport but observed that Mapuche

families transport 100 kg of seeds per trip. Given the large size of the Araucaria cone, it

was almost certainly discarded.

Pine nuts

For Portugal, the charcoal data are equivocal concerning the types of pine located

near the archaeological sites of the Bølling/Allerød phase. Figuieral (1993, 1995) has

developed some criteria to discern Pinus pinaster from P. pinea. Distinguishing these two

is important because P. pinaster has a much smaller seed than the ‘pignolia’ of P. pinea.

Initial research led Figueiral (1993) to argue that both types were present in the charcoal

assemblage from CPM. Subsequent SEM analyses may suggest that much of the pine

charcoal from CPM was from P. pinaster but the assemblage was not systematically

reanalyzed (Figueiral 1995). The conclusion is based solely on impressions and the

methodology itself is new and so far has not been replicated by others. More SEM analyses

are needed in order to verify these criteria. Considerable variability and overlap in the

criteria offered by Jacquiot, Schweingruber and Figueiral makes it difficult to accept that

these pines can be reliably differentiated by their wood anatomy at present. In evolutionary

terms, the wood is under the least amount of selection and thus inter-species variation is

low. Most species within a given genus are only distinguishable by leaf structure and

form, flower and seed morphology

Considering the preferred habitats of these pines, the calcareous soils of the limestone

uplands in Estremadura may have limited the spread of P. pinea. Thus, sites in Serras of

Aire and Candeeiros, where most of the fauna-bearing caves are located, would not be

expected to have evidence of pine nut consumption. As will be seen in the next chapter,

these sites are specialized animal carcass processing locations, not residential sites (Bicho

1996; Zilhão 1995). On the other hand, the low plains and valleys of Estremadura are

covered by Miocene sands and podzols which are well suited for P. pinea. The Rio Maior

valley would have been prime habitat for P. pinea. Its occurrence during the Tardiglacial

is confirmed by the charcoal record. Unfortunately, direct evidence of pine nut processing

in open-air sites is missing. Organic preservation is extremely poor in the sediments of

the area. The absence of charred nutshells does not necessarily mean that pine nuts were

not utilized. Although heating aids seed removal and thus may result in charring, it is not

required. Dried seeds could easily be cracked open to remove the seed from the hull. The

technology does not necessitate large slab grinding stones, common for plant processing

in later times. Small grinding stones have been found at CPM in the Rio Maior valley but

no residue analyses have been made on them. Grinding stones were also found in open-

air Gravettian sites in the Rio Maior valley and Vale Boi in southern Portugal (Thacker

pers. com; Bicho et al. 2000). Of course, the occurrence of grinding stones does not

necessarily mean they were used in plant processing. They could have been used in

pigment grinding or animal bone grease processing. Until residue analyses are made,

interpretation of their use is open.

For Mediterranean Spain, Badal (1998, 2001) identified over 9,900 carbonized pieces

of pine cones and seed hulls from Pinus pinea in the Cueva de Nerja. Though natural fires

can ignite vegetation in caves, the association of these pieces with archaeological

occupations provides a reasonable basis for considering them human food refuse. The

majority were collected in three archaeological levels dated to the Solutrean, Upper

Magdalenian and Microlaminar Epipaleolithic. In the Solutrean level 9 dated 18,420+/-

530 bp to 17,940+/-200 bp (Jordá Pardo et al. 1990), Badal (1998) reported 2580 fragments

of cones and 196 hull pieces. The occurrence of P. pinea indicates warm, humid conditions

near the cave suggesting an occupation during an interstadial following the LGM.

Importantly, it demonstrates substantial human use of pine nuts as early as the Solutrean

in the cave, although pine cones and hulls are found in the Early Upper Paleolithic levels

too. Equal frequency of pine nuts occurs in the Upper Magdalenian level 7 dated 12,130

+/-130 bp corresponding to the Bølling/Allerød interstadials. Large numbers of cone

and hull fragments are also found in the subsequent Microlaminar Epipaleolithic level

dated 10,860+/-160 bp. While this date falls within the Younger Dryas period, it overlaps

the end of the warmer Allerød. Based on faunal analyses, the Younger Dryas at Nerja is

characterized by cold-adapted birds and fish so it is unlikely that vegetation remained the

same. Therefore, the pine nuts in the Epipaleolithic levels were likely deposited prior to

the Younger Dryas.

In all levels containing evidence for pine nuts, the occurrence of cone fragments implies

a short distance to pine stands. According to the Barlow and Metcalfe (1996) model, these

would only be transported with pine nuts if round trip times to find and collect them

were less than a couple of hours.


According to pollen and charcoal analyses, both evergreen and deciduous oaks were

present in Late Pleistocene Portugal. Generally, only broad categories may be discernible

by wood anatomy. Oaks are divided by non-taxonomic categories, such as white oaks,

black oaks red oaks, etc., by North American foresters. Identification through wood

anatomy is more easily done for these categories than by species although there is apparent

disagreement on this between North American and European specialists. Carrión et al.

(2000) claim to have distinguished the pollen form of Q. suber from Q. ilex. Their criteria

have not been applied to other regional pollen analyses so the best hope for determining

which species was more prevalent during the Bølling/ Allerød phase is the modern ecology

and biogeography of the two species. Regardless of these problems, each of the species

present today would have been present in the Late Pleistocene. Because Iberia was a

refuge for many northern European plants, additional deciduous types may be present in

the pollen and charcoal samples from northern Iberian sites (Bennett et al. 1991).

During the cold periods of the LGM and Dryas I, Q. ilex was probably limited in its

biogeographic range. While it may have been able to withstand the lower temperatures,

the increased aridity would have precluded the formation of closed-canopy Holm oak

forests that characterize much of the natural areas of lowland central and southern Portugal

today. Charcoal analyses of LGM sites shows that Q. suber was present in Caldeirão,

located in a sheltered valley, but not in the upper elevation site of Anecrial or the open,

low-elevation site of CPM (Zilhão 1997). Thus, acorns from these species would not have

been available in large quantities to hunter-gatherers during cold, dry periods like the

Solutrean and Middle (?) Magdalenian. They would likely have been present during the

immediate post-LGM Early Magdalenian but more prevalent during the warm, humid

Bølling/Allerød period corresponding to the Late Magdalenian. The evergreen oak

charcoal from CPM attests to its presence in the Late Magdalenian. The occurrence of Q.

suber in CPM and Caldeirão is likely due to their location near sandy Miocene sediments

where cork oaks are found today.

At present there is no physical evidence of acorn use during the Upper Paleolithic or

Epipaleolithic in central Portugal. In Spain, acorn parts were identified in the Upper

Paleolithic levels of Cueva de Nerja and a few Epipaleolithic sites in Catalunya mentioned

earlier. Considering the Bettinger et al. (1997) model for acorn processing and transport,

unprocessed acorns should be transported from stands to residential sites. They found
Table : Comparative nutritional values expressed in g/g; mg/g and µg/g for
selected nuts from Mediterranean woodlands

Pinus Pinus Pinus Quercus Quercus Castanea Corylus

pinea pinea pinea ilex suber sativa avellana






Miner al s
Vitam ins

Data from USDA food composition database  Alce Ingeniería (Spain) Tabela da Composição dos
Alimentos Portugueses (Gonçalves Ferreira & da Silva Graça  )

Table : Comparative nutritional values expressed in

g/g; mg/g and µg/g for selected birds
from Mediterranean wetlands/woodlands
Common name Duck Duck Partridge
Portuguese name Pato Perdiz


Minerals (mg)




Folate  µg  
A (IU)  
B  µg  
Data from USDA food composition database ; Alce Ingeniería (Spain)
Table  : Comparative nutritional values expressed in g/g; mg/g and µg/g for
selected game from Mediterranean woodlands
Common Red deer Red deer Roe deer Wild boar Wild boar Rabbit Rabbit Goat Goat Horse Horse
Portuguese Veado Corço Javali Coelho Cabra Cavalo
Minerals (mg)




Folate     µg  µg  µg  µg   
A (IU)           
B   µg  µg 
µg  µg  µg  µg  µg  µg  µg  µg
Data from USDA food composition database ; Alce Ingeniería (Spain)

Table  : Comparative nutritional values expressed in g/g; mg/g and µg/g
for selected fish and from Iberian waters
Common name Salmon Sardine  Gilthead Shad Trout
Portuguese name Salmão Sardinha Dourada Sável Truta

(   ) (

Minerals (mg)
A (IU)     
B  µg  µg µ   µg
Data from Nettleton ( ) Tables  &  ; Alce Ingeniería (Spain);
Tabela da Composição dos Alimentos Portugueses (Gonçalves Ferriera & da Silva Graça  )
All analyses conducted on fresh fish
Table : Comparative nutritional values expressed in g/g; mg/g and µg/g
for selected shellfish and from Iberian waters

Common name Cockle Clam Razor clam Mussel  Oyster Scallop Limpet Periwinkle
Portuguese name Berbigão Amêijoa Navalha Meixilhão Ostra Vieira Lapa Burrié


(  ) (  ) ( ) ( )



Minerals (mg)

Folate      µg   
A (IU)        
B µg    µg  µg   
Data from Nettleton ( ) Tables  &  ; Alce Ingeniería (Spain); Tabela da Composição dos
Alimentos Portugueses
(Gonçalves Ferriera & da Silva Graça  )
All analyses conducted on fresh shellfish

that cupule removal, cracking, winnowing and leaching or roasting are not economical in

the field unless acorns are transported over 125 km. Since no groups were ever observed

transporting acorns over such long distances it is assumed that this processing did not

take place in the field. Drying to reduce load weight may be the only economical field

processing but this is contingent on the amount of time spent in the field. Therefore,

evidence of acorn use should occur predominately in residential camps, not in logistical

task sites. Unfortunately, these are the sites in Portugal that lack organic preservation.

Nutritional utility

As discussed in chapter 2, the evolutionary ecological studies of tropical foragers

have shown that reproductive fitness, not energy maximization, is the primary factor

driving subsistence strategies (Hawkes et al. 1991, 1997, 2001; O’Connell 2000; Winterhalder

and Smith 2002). The nutritional ecology approach offers an additional perspective to

dietary choice by considering the health and reproductive consequences.

Tables 4.10, 4.11, 4.12, 4.13 and 4.14 show the nutritional values of food types that

were available to Late Pleistocene hunter-gatherers in central Portugal. Note that small

game like rabbits offer the same proportions of protein, fat and micronutrients as large

game if not higher. Birds are also high in protein and much higher in fat than terrestrial

mammals. Fish are also comparable sources of protein. Shellfish contain lesser quantities

of protein the proportions of various proteins are better suited to human metabolism (Wing

and Brown 1979; Nettleton 1985; Erlandson 1988). Plants on the other hand have a wide

range of nutrient values but are the best source of carbohydrate. In the case of tree nuts,

pine nuts and hazelnuts have high proportions of protein and fat and, consequently,

immensely high caloric value compared to animals. Acorns and chestnuts are generally

good sources of carbohydrate. Detailed nutritional analyses of other edible wild plants

such as seeds, leaves, roots, bulbs and tubers are not available. However, some qualitative

information is provided in Table 4.2. For the most part, these would have provided

vitamins, minerals, fiber, essential oils and fatty acids. These types of plants would have

been better sources of vitamins C & E than animal organ meat.

While post-encounter return rates for all the resources available to Upper Paleolithic

people in Iberia are not feasible, the conclusion reached by many that large game will

always outrank small game, aquatic resources, and plants in terms of caloric energy is a

useful starting point for discussing foraging efficiency in energetic terms (Kelly 1995;

Broughton 1995; Broughton and O’Connell 1999; Ugan and Bright 2001). Assuming the

larger the animal, the higher the rank, horse and aurochs should be the highest ranking

resource since they would arguably provide the greatest number of calories per animal

unit even though they might not be encountered frequently. Medium ungulates like red

deer, ibex, chamois and wild boar might be roughly equivalent in caloric value but much

lower than large animals and encountered more often. Small game such as rabbits and

birds would rank lower due to their smaller package size and despite their greater

abundance and encounter frequency. Plants and shellfish, while highly productive, would

rank lowest because they require higher labor investment for much lower caloric return

as Bailey and others have argued.

However, resource rank does not often predict the relative contribution to the diet. If

generalizations about modern hunter-gatherers can be extrapolated to the past, then


perhaps large game hunting was driven more by male status-seeking than subsistence

concerns (O’Connell 2000). In places like the temperate mid-latitudes and sub-humid

Mediterranean where plants and small animals were always present, large game hunting

might not have been as important to overall subsistence as in the polar regions where

there is little else to eat. Daily subsistence may therefore have been based on plant collection

and small animal procurement by all segments of the population.

On the other hand, if Jochim’s (1998) claim that hunter-gatherers will naturally choose

a balanced diet is a result of our evolutionary heritage as primates then meat would not be

the central focus of prehistoric diets given the problems discussed in chapter 2 (Speth and

Spielmann 1983; Eaton et al. 1997, 1998, 2002; Cordain et al. 2000a&b, 2002). Hockett and

Haws (in press) have argued that balanced essential nutrient intake through dietary

diversity enabled Early Upper Paleolithic humans to grow their populations and replace

archaic forms in Europe. In Iberia, diverse diets may have a greater antiquity due to

greater availability of plants and small animals. Given this it is more plausible to assume

that modern humans in Iberia were naturally acquiring a balanced diet during the Upper

Paleolithic of Iberia through dietary diversification.

In nutritional terms, the values for the foods in Tables 4.10- 4.14 show that plants and

small animals like shellfish offer many nutrients in better proportion than ungulates.

Hunter-gatherers would have achieved better overall health and greater reproductive

success by exploiting a wider range of resources. Nutritionally, tree nuts may have been

important substitutes for meat in lean seasons due to their storability. In the Mediterranean

region, including Portugal, the lean season would have been the summer dry season.

Pine nuts would have maintained their fat stores for several months enabling hunter-

gatherers to offset the risk of protein poisoning if they were eating large amounts of meat

in the absence of plants. They could also have complemented meat during seasons where

both were plentiful. Stores of carbohydrates in the form of acorns and/or chestnuts would

also benefit people through their protein-sparing action. The leafy greens with their

essential vitamins and carotenoids would have been available for much of the year due to

the mild, rainy winters.

4.5 Discussion

Where edible plants are found people will consume them. This is obvious from

ethnographic observations and global surveys of hunter-gatherers. The appearance of

edible portions of various plants in the Early Holocene sites of Catalunya, Andorra and

southern France tracks the changes in the environment. In these regions climate was

more rigorous during the Late Pleistocene as evident by charcoal analyses, pollen and

deep sea cores in the northwest Mediterranean. As forests spread so does evidence of the

use of their products. In areas such as southern Spain and central Portugal where climate

was less severe the pollen and charcoal record shows that the types of plants exploited in

later times were present in the same proportion as today by 12,500 bp. If the hypothesis

that humans will consume plants when they are available is true, then there should be

evidence of their use during this period. Sites in southern Spain do contain evidence of

plant exploitation before the Early Holocene. However, there is no such evidence of

macrobotanical remains in central Portugal. This is largely due to excavation methods


and the non-use of techniques suited to recovering plant remains. Most sites were excavated

before modern techniques like flotation were developed. Research bias against the idea

that plants were important components of Paleolithic diets also made attempts at recovery

seem pointless to previous excavators.

Using the expectations derived from analogous species of pine nuts and acorns in the

Great Basin and California, there would be little reason to expect either to have been

processed in Lapa do Picareiro, Caldeirão, Lapa dos Coelhos, Bocas or Lapa do Suão.

Neither require intensive field processing with the exception of removing pine seed hulls

from their cones. These would likely have been deposited in locations within pine stands

or in residential sites, not specialized animal carcass processing sites. Acorns would only

be processed and inedible fractions discarded in residential sites as well. No features

have been identified as leaching pits in the open-air Upper Paleolithic, Epipaleolithic or

Mesolithic sites of central Portugal. Charred nutshells of either pine nuts or acorns have

not been recovered. No site has been excavated using flotation or any other systematic

method for recovering these items. Recently, wet-sieving has been done using bulk samples

from a few sites but this has not been done for or by paleoethnobotanists. All charred

material is treated as charcoal and only recently has identification of wood species occurred

prior to sample destruction for radiocarbon dating (Figueiral 1993). No report on charcoal

remains has mentioned plant parts other than wood.

It can be demonstrated that central Portugal was a refugium for Mediterranean and

temperate plant species during the Late Glacial. This mixed community existed in low-

lying plains and valleys until the Late Glacial interstadials allowed recolonization of upper

elevations. However, the archaeological record is insufficient to know whether or not

plants were a substantial portion of the diet during the Late Upper Paleolithic and

Epipaleolithic. Current research projects in central and southern Portugal are utilizing

methods to recover such evidence. Preliminary data indicate the use of plants as early as

the Gravettian (Thacker pers. com.). Given these findings, it is likely that plant exploitation

was a regular and probably significant portion of the human diet during the Late Upper

Paleolithic. The implications from central place modeling drawn upon in this chapter

should serve to guide future research expectations concerning the types of plant materials

that could be expected occur in a given location.

Based on energy-considerations alone, the central place models applied to analogous

resources shows that certain plants can be economically exploited over long distances.

The Mapuche example cited above further illustrates the value of plants with regard to

macronutrient content. The nutrient values for the tree nuts of the Mediterranean are

comparable and/or complementary to those of animal resources. If people were under

resource stress, the prime motivation for subsistence change in the Broad Spectrum

Revolution model and diet breadth model, it is doubtful that people would have ignored

such an abundant and easily processed resource such as pine nuts. The labor inputs and

nutritional gains, whether energetic or marcronutrient, would have made pine nuts as

highly ranked and probably higher than those from the Great Basin. Although late fall

and winter would have been ideal collection times, immediate consumption would not

have been necessary. Cones or seeds in hulls could be stored for several months, increasing

their value for human diets.


Pine nut and acorns serve as examples of the types of plants that could have played

important roles in Upper Paleolithic hunter-gatherer subsistence in Portugal. Whether or

not they could have been staples is debatable. Additional nuts may not have been as

abundant as pine nuts and acorns in Estremadura. Walnuts are thought to have been

introduced to Portugal by the Romans. Chestnuts and hazelnuts occur naturally and

were evident from Late Pleistocene pollen diagrams in northwest Iberia. However, neither

have been identified in charcoal assemblages from Estremadura.

Certainly, a wide variety of plants besides tree nuts were available as food. Many of

these, such as greens and tubers leave no durable remains. The pit of wild fruits and

berries are virtually unknown in archaeological sites from this period. Only a few seeds

from Rubus, probably blackberry, were recovered at Picareiro. Of the edible roots, tubers

and bulbs mentioned by Clarke (1976), there is no specific evidence to date. However, the

use-wear and starch grain analyses on Gravettian tools has produced some promising

results (Thacker pers. com.). The availability of these plants is unknown because there are

no pollen cores in low altitude locations in central Portugal that date to the Late Glacial.

However, the pollen diagram from San Rafael in Spain shows that coastal marsh taxa

were present as early as 16,000 bp. Even at Mougás, in Galicia, pollen from the Liliaceae

and Umbelliferae, two important families with edible taxa, were present in the Bølling/

Allerød. Since many of the plants Clarke discussed inhabit this zone, they may have been

available to Magdalenian people. The lack of human skeletal material on which stable

isotope and trace element assays can be made precludes a better understanding of the

sources of protein and calcium in the diet and the role of marine foods.

In the next chapter, the representativeness of animal resources in the Late Pleistocene

and Early Holocene of central Portugal is addressed. Archaeological evidence for animal

exploitation is investigated through a consideration of prey behavioral ecology and

taxonomic identification. Site function and formation processes are investigated through

taphonomy and skeletal element representation. This enables an understanding of the

nature of animal exploitation and the role of animals in prehistoric diet during the Late

Upper Paleolithic in central Portugal.


Chapter 5: Upper Paleolithic faunal exploitation in central Portugal

5.1 History of investigation

Pleistocene faunas

The study of the Paleolithic in Portugal began in the 19th century (Bicho 1992, 1993;

Zilhão n.d., 1995). The initial goal of early studies was to confirm the coexistence of humans

and extinct Pleistocene animals in Portugal. J.F. Nery Delgado and Carlos Ribeiro, of the

Serviços Geológicos, were aware of work in northern France proving the antiquity of man.

Excavations by Delgado at the Gruta de Casa da Moura, just six years after the finds of

Boucher de Perthes in the Somme Valley were accepted, showed this to be true in Portugal

as well. His work was based on a consideration of the agents responsible for the animal

bone deposition and the formation processed of the cave deposits.

By the early 20th century, taphonomic considerations appear to have been forgotten

or ignored. Vieira Natividade and others from the Serviços Geológicos excavated numerous

fauna-bearing caves in the area between the Estremadura coast and the limestone massif.

Many caves were excavated without regard for taphonomy or stratigraphy as was common

in many regions during this period. Of these, Gruta das Alcobertas and Algar de João

Ramos contained Pleistocene fauna (Araújo and Zilhão 1991; Cardoso 1996). The former

has an undated Upper Paleolithic occupation while the latter is considered a paleontological

site which was recently dated to 14,170 +/-170 bp (Antunes 1989). Edouard Harlé (1910-

11) published an extensive paper on Pleistocene mammals in Portugal, which summarized

the fauna collected from all of the known Pleistocene sites. His work remained the primary

source on the subject until the 1970s. During the interim, Quaternary research flourished

under Henri Breuil, Georges Zbyszewski and Octávio da Veiga Ferreira of the Serviços

Geológicos but few Late Upper Paleolithic sites were excavated. Most of their research

focused on the period between the Last Interglacial and Last Glacial Maximum. Jean

Roche (1972) integrated the geologic work with a summary of Pleistocene faunas. This

was a descriptive paper that used the faunal record as a guide for estimating glacial climate

conditions. Recently, Cardoso (1995) systematically reanalyzed the large mammal

collections in the Serviços Geológicos from a paleontological perspective. His compendium

of fauna and the sites from which they came is a useful guide for comparative biometrics

but offers little in the way of understanding human behavior. No studies of prey mortality

profiles, skeletal element representation, or butchery patterns have been made on these

materials. In fact, the nature of excavation methodology and museum curation precludes

these analyses since the assemblages represent only a partial sample of the original site

contents. Most of the sites are caves that date to the Middle Paleolithic or Early Upper

Paleolithic and contained assemblages accumulated by humans and carnivores. Each are

characterized by the presence of numerous large and medium carnivores such as hyena,

cave lion, leopard, lynx, bear, wolf, cuon and fox. Herbivores include horse, aurochs, red

deer, roe deer, ibex, chamois, wild boar with some elephant, rhinoceros, and hippopotamus

prior to OIS 2 (Cardoso 1997). Hedgehogs, birds, tortoises, land snails, fish and/or shellfish

are listed in most sites but their presence is difficult to evaluate in the absence of taphonomic


Late Upper Paleolithic faunal exploitation

On the Iberian Peninsula, rabbits have been hunted for tens of thousands of years. In

fact, the name ‘Hispania’ or ‘España’, derived from the Phoenician word ‘span’, means

‘land of rabbits’ (Bocharti 1712). In Mediterranean Spain and Portugal, Upper Paleolithic

cave and rockshelter sites are usually full of rabbit bones. In Spain, researchers have long

held that humans deposited the large numbers of rabbit bones found in Upper Paleolithic

cave and rockshelter sites. Most researchers viewed rabbit as completely insignificant in

dietary terms despite their high frequencies in Paleolithic sites. In some cases, rabbit

bones were not kept by the excavator (e.g., Pericot at Parpalló). Estimations of dietary

contribution based on meat weight has left rabbit completely marginalized by most

archaeologists who have worked in Iberia. In Portugal, the first consideration of rabbit

taphonomy was Delgado’s (1867) study of Casa da Moura. The earliest description of

Pleistocene faunas listed “nombreux restes” of rabbits at Algar do João Ramos and Casa

da Moura (Harlé 1910/11). The latter was reported to have “rabbit bones layers” that

Breuil (1918) subsequently identified as Magdalenian. These materials were unfortunately

destroyed by fire in 1975. Systematic taphonomic studies of raptor, carnivore and human

damage to rabbit bones only began in the last 15 years with the work of Hockett, Perez-

Ripoll and Sanchis Serra (Hockett 1989, 1991, 1992, 1994, 1995, 1996, 1999; Pérez-Ripoll

1993; Rowley-Conwy 1992; Sanchis Serra 2000).

For the Late Upper Paleolithic the first site excavated with careful attention to

recovering faunal remains was Lapa do Suão or Suão Cave. Prior to this only Casa da

Moura and Bocas rockshelter, excavated in the 1930s, had indications of Magdalenian

subsistence. The Bocas material is housed in the National Museum and is being studied

by Maria João Valente. This site is discussed below. Since the late 1970s and early 1980s,

the fauna from Suão served as the basis for interpreting Magdalenian subsistence in

Portugal (Roche 1979). Yet, strangely the assemblage was never analyzed and the only

information published was a list of the identified species, with the exception of the single

numerically dominate species, the European rabbit. Consensus was that Magdalenian

subsistence was characterized by a focus on red deer, wild caprids, horse, aurochs and

wild boar (Bicho 1993, 1994; Marks and Mishoe 1997; Roche 1979; Zilhão 1992, 1997).

Rabbits were not recognized as a significant economic resource until recently. The emphasis

on large game has a long tradition in European Paleolithic subsistence studies and this

perspective was deeply ingrained in Portuguese archaeology as well.

The excavation of Gruta do Caldeirão in the 1980s by Zilhão added a significant,

deeply stratified multi-component sequence with faunal remains from the Neolithic to

the Middle Paleolithic. This record confirmed the general picture of Upper Paleolithic

subsistence with a further refinement of the Solutrean and Magdalenian. Zilhão (1992,

1997) argued that Solutrean subsistence was largely focused on horse, red deer, ibex and

chamois but that Late Pleistocene climate change eliminated ibex and chamois resulting

in a Magdalenian subsistence based largely on red deer and rabbit (see chapter 3).

To date, the most important Late Upper Paleolithic site with preserved faunal remains

is Lapa do Picareiro. This multi-component site represents the most fine-grained evidence

for Magdalenian subsistence in Portugal. It contains several radiocarbon-dated levels

occupied between 8,000 and 12,500 bp. The deposits continue well below but are not well

dated. The three sites and their Magdalenian levels are the focus of this chapter.

5.2 Prey behavioral ecology

Modern prey population structure and reproductive and behavioral patterns have

been used for many years to interpret prehistoric hunting strategies. The animals hunted

in Portugal during the Late Pleistocene (after the LGM) are still present today in Iberia

with the exception of auroch and wild horse. Aurochs were hunted until at least the

Copper Age and perhaps into Roman times (Estévez and Saña 1999). Horse was hunted

by Magdalenian people but its wild ancestors disappeared from Iberia in Late Prehistory.

The relationship between the Iberian horse and modern domestic ones is subject of debate

(e.g., Uerpmann 1996). Ibex, extirpated in Portugal during the 19th century, and chamois

still live in protected areas in Spain. Red deer, roe deer, wild boar, rabbits and birds still

live in Portugal today. The carnivores wolf, fox, lynx , badger and otter are present in

very low numbers as their prey and habitat have been reduced for centuries. Of these,

only fox is known from Picareiro while wolf, lynx and badger are known from other

contemporary sites such as Suão and Caldeirão. Otter has not been identified in Upper

Paleolithic sites but that may be due to site location.

Because present habitats are substantially different than those of the Late Pleistocene,

little can be said of the animal biomass potential without making some speculative leaps.

The fact that many of the animals hunted in the Late Pleistocene are still present points to

their remarkable adaptability in the face of climatic and environmental change. Despite

these changes and the fact that past distributions may have been different, the modern

ecology is considered here using uniformitarian principles in order to understand the


structure of the archaeofaunal record. Given the variability in social and feeding behaviors

observed within ungulate species occupying different habitats, modern studies should

not be used as rigid guidelines for interpreting the past. A brief discussion of each of the

economically important animal species follows.

Red deer, Cervus elaphus

Red deer in Iberia prefer open forests of oak. Comparative studies suggest higher

population densities for red deer in open versus closed forests (Straus 1981). Their diet is

mainly leaves, shoots and twigs of trees and shrubs but they will also eat acorns, chestnuts

and grasses. During summer, when rainfall is low they may inhabit piney areas. Given

the open-forested nature of Late Pleistocene Iberia it is not surprising that red deer is the

most abundant ungulate in many archaeological sites.

Male red deer can reach 180 kg while females are usually less than 100 kg. Males

shed their antler between February and April and they regrow around July or August

(Mathias et al. 1998). During most of the year males are solitary while females maintain

small groups of juveniles born the previous spring with a few 1-2 year old animals. Red

deer do not migrate long distances. Males usually occupy territories of 6-10 km2 while

females have ones ranging 2.5-8 km2 (Soriguer et al. 1994). Rutting season occurs during

fall. Males form harems that average about 10 animals. This represents the best time to

take large numbers of red deer in a single hunt. In spring, females break away to give

birth and rear young. Today this occurs between April and June in Portugal and May and

July in Andalucia (Mathias et al. 1998; Soriguer et al. 1994). Juveniles develop rapidly

achieving half the body weight of an adult by the first year. Growth slows afterwards

with females reaching maximum size around 4-5 years and males around 7-8 years (Mathias

et al. 1998).

Wild Boar, Sus scrofa

Wild boar are found in deciduous and evergreen forests and dense, low shrubland.

They are omnivores and eat mainly fruits, nuts, tubers and roots, but also insects, snails,

frogs, lizards, rodents, eggs and even dead birds (Mathias et al. 1998; Serôdio 1997). During

the fall and winter they fatten up on chestnuts and acorns which together can make up

about 60% of their diet. Wild boar never stray far from permanent water sources but will

walk several kilometers to find chestnuts and acorns (Serôdio 1997). Adult males can

reach 100 kg on average. Females are slightly smaller at around 80 kg. Generally, adult

males are solitary are stay in small groups of two to three during the year until the

reproductive season. Females tend to lead small groups of juveniles. Mating occurs

between November and April. After a gestation of 4 months, births usually range from 2-

4 animals but as many as seven are possible. Some females have been observed giving

birth twice in one year; once in January-February the second in August-September.

Whether or not this occurred in prehistory is unknown, but it raises potential problems in

seasonality determinations based on tooth eruption or epiphyseal fusion.

Auroch, Bos primigenius

Little is known about the lifeways of the auroch. The last one was killed in the

early 17th century in eastern Europe. Based on biometric data, adult male aurochs are

thought to have reached about 800 kg and heights of 1.5- 2 m (Guintard 1999). Biometrics

have also confirmed sexual dimorphism. Males are thought to have been solitary since

many isolated male aurochs skeletons have been found in bogs across northern Europe.

Some of these were wounded by hunters as in the case of the well known Prejlerup aurochs

that had associated microliths from 9-15 arrows (Aaris-Sørensen 1999). In Iberia, aurochs

were generally larger in the south than in the north, possibly due to the milder and more

humid climate (Estévez and Saña 1999). The aurochs may have been a very important

animal for Late Upper Paleolithic people as it is one of the best represented animals in the

Upper Paleolithic rock art of the Côa Valley in northwestern Portugal. At Domingo García

and Siega Verde in central Spain, La Pileta in southern Spain and Parpalló in eastern

Spain, images of aurochs were also a substantial proportion of the total number (Weniger


Ibex, Capra pyrenaica

Ibex still lives in isolated mountain ranges all over Spain. They are gregarious and

prefer open areas above ~600m elevation. However, they easily adapt to more forested

areas of Mediterranean woodlands (Garcia-Gonzalez and Cuartas 1992). In the Sierras de

Cazorla and Segura of southern Spain, ibex are found between 1300m and 2000m (Alados

1985). Mature males live in hierarchical groups of 20 or more animals. During the year

females form groups of mixed juveniles. In May when birthing begins, females rear the

newborns while 1-2 year-old males break away and form smaller groups. In August,

these animals rejoin the female groups. The fall rut occurs in November and December

when prime-aged males drive away older and sub-adult males to join with females (Alados

1985). In January, the groups segregate again until the following autumn (Alados 1985).

Additional studies show that ibex move seasonally through altitudinal zones. During

winter ibex descend to low altitude woodlands and open valleys. Males may occupy

woodlands more than females (Alados 1985). During warmer months when males gather

increasing numbers, they ascend to shrubland pastures. In summer, the males are found

on steep slopes. This group dynamic probably relates to ibex forage quality, the ideal

being more shrubland type. In the Sierra de Cazorla, population densities are the highest

in elevations 1300-1400m. In this zone ibex is the most abundant ungulate and their diet

is also the most diverse, though the preference is for leaves and branches of Quercus ilex,

the holm oak (Garcia-Gonzalez and Cuartas 1992). Increased dietary diversity in the lower

altitudes is thought to result from competition with other ungulate species. “The increase

in diet diversity values in May could be explained by a wider availability of species and

by a nutrient balancing strategy” (Garcia-Gonzalez and Cuartas 1992: 169). This last

statement may have important ramifications for energy-based optimal foraging models.

Chamois, Rupicapra rupicapra

As mentioned in the previous chapter, chamois has been used as a paleoclimatic

indicator for cold dry conditions during the Last Glacial Maximum due to its greater

frequency in southern France during cold periods (Delpech 1978). Contrary to many

suggestions, chamois and ibex should not be used together to indicate certain

paleoenvironmental conditions. Though modern data show that chamois typically live in

temperate montane areas of fairly low precipitation, their past distribution may have been

quite different (Miracle and Sturdy 1991). In Herzegovina, Miracle and Sturdy (1991)

found chamois bones in Late Upper Paleolithic sites that were located in paleoenvironments

similar to modern low elevation zones that chamois do not occupy today. This suggests

that chamois had a much wider distribution in the past. Tosi et al. (1987) have observed

chamois in protected zones in Italy regularly occupying chestnut and walnut forests as

low as 300m asl. Chamois exhibit a similar group structure to ibex in that the females

form large groups (over 100 animals) during spring and summer, while males form slightly

smaller ones (Berducou and Bousses 1985). The rut occurs in November and December.

Females give birth in June. In winter, chamois inhabit shady, forested areas while in

summer they ascend to higher altitude zones. In Spain today they migrate seasonally 600-

900m between summer and winter ranges. Territory is greatly restricted in winter

compared to ibex and wild sheep (Garcia-Gonzalez et al. 1992).

Horse, Equus sp.

There are no living analogs for wild equid behavioral ecology in southern Europe.

Two distinct species of horse occupied Iberia during the Late Pleistocene: Equus hydruntinus

and Equus caballus (Cardoso 1995). Like its modern representatives horses are grazing

animals occupying grassy plains. In Estremadura, they are rare during the Late Glacial

and Early Postglacial, probably due to site location and increased forestation.

Rabbit, Oryctolagus cuniculus


The European rabbit probably evolved on the Iberia Peninsula. These animals are

very abundant and territorial. They live primarily in burrows in densely packed warrens.

Rabbits are quite fecund, giving birth throughout the year with spikes in birth frequency

during rainy periods in Spain and Portugal (Garson 1981; Soriguer and Rogers 1981).

Therefore rabbits would have been regularly available throughout the year. Average size

is about 1-2 kg for an adult rabbit. With large numbers of rabbits living in restricted

territories, prehistoric hunters would have been able to easily collect sufficient quantities

to make rabbit a significant dietary component.

With the behavioral ecology of Late Pleistocene prey in mind, the Late Upper

Paleolithic archaeofaunas can be interpreted. Attention now turns to the sites of Lapa do

Picareiro and Lapa do Suão, which contain the most reliable stratigraphic contexts for

Magdalenian subsistence. These sites are then compared to Caldeirão and other sites in

the region.

5.3 Lapa do Picareiro

Lapa do Picareiro is a shallow cave located near the town of Covão do Coelho in a

small depression on the west side of the Serra d’Aire at about 500m above sea level (Figure

5.1-5.4). Gil Andrade first investigated it during the 1960s, when several human burials

associated with decorated pottery were found. The name ‘lapa’ was given to refer to the

shape of the cave. ‘Lapa’ means limpet in Portuguese and the cave is shaped like the

inside of a limpet shell. The site was ignored until 1988, when João Zilhão and members

of STEA (Sociedade Torrejano de Espeleologia e Arqueologia) revisited it. They recovered


Figure 5.2: Picareiro is at the top of the Serra d’Aire (above) and 5.3: entrance
Figure : Plan of Lapa do Picareiro




Cave profile


Figure 5.5: Cave prior to excavation (above) and 5.6: after 2001 (below)

charcoal, flints, and animal bone suggesting an Upper Paleolithic component to the site.

In 1994, Bicho tested the site as part of a project to record all of the archaeological sites in

the region (Figure 5.5). This testing not only confirmed the suspicion that the site was

occupied during the Upper Paleolithic, but that it had at least two occupations, the

uppermost dated to 10,000 bp, and that the deposits continued deeper. As a result, Bicho

systematically excavated the site between 1995 and 2001 (Figure 5.6). Beginning in 1995,

visible archaeological materials were piece-plotted and the sediment fine-screened, washed

and sorted to recover small rodent bones, fish vertebrae, shell, charcoal, and numerous

backed bladelets and microdebitage (Bicho et al. 2000).

Table : Radiocarbon dates for Lapa do Picareiro

Stratum Sample Date bp* Lab 

D Charcoal    Wk

E Upper Charcoal 


E Middle Charcoal 


E Lower Charcoal   Wk 

F Charcoal 

F Charcoal    Wk

G Charcoal     OxA

* noncalibrated results

The excavation revealed 19 stratigraphic levels labeled A-S, with the 6 dated

archaeological occupations occurring in D, E, F, and G (Figures 5.7 & 5.8). Levels I and J

have archaeological deposits but they are undated. Of the dated levels, D dates to 8,300, E

Figures 5.7 & 5.8: Stratigraphic profile of Lapa do Picareiro


Upper to 10,070 bp, E Middle to 11,700 bp and E Lower to 11,500 each with a standard

deviation of 120 years. Level F dates between 11,700 bp and 12,300 bp, while G dates to

12,300 bp. The large mammal remains from these six, plus I and J, recovered during the

1995-2000 excavations were analyzed and the results are presented here.

Figure 5.4 shows the site plan with the excavated area. Each 1 m2 unit was excavated

to Level K, an undated Early Upper Paleolithic level with few lithics and faunal remains.

The faunal sample discussed below is entirely Magdalenian and represents about one-

third of the cave deposits. The assemblages from the earlier levels are extremely sparse

and are not adequate to consider long-term diachronic changes in subsistence. The

Magdalenian ones are sufficient to discuss the nature of site function during the

occupations. However, they probably do not represent the full range of subsistence

behaviors that people engaged in during the Magdalenian.

Faunal analysis for Lapa do Picareiro

During the excavations, several thousand bones were piece-plotted and many more

recovered through screening. Fauna include red deer, roe deer, wild boar, auroch, chamois,

rabbit, rodents, marine shellfish (mussels, limpets, and clams), sardine, and land snails.

The assemblage is dominated by rabbit bones, over ten thousand of which have been

studied by Bryan Hockett (BLM). There are a total of 1,837 medium to large mammal

bone specimens in the assemblage. Of these, 451 (25%) can be identified to specific taxa

(Table 5.2: NISP). After rabbit, the primary species in the assemblage is red deer, which

represents 65% of the identified macrofaunal assemblage. This is followed by wild boar

at 30%, aurochs at 3%, and chamois and ibex each less than one percent. The rest are

indeterminate but can be divided into general size classes. Of the 1,360 specimens, 408

(30%) are Cervus-sized while the remaining 70% can be classified as small-medium

ungulate. This latter category probably includes bones from wild boar, caprids and

subadult red deer. However the pieces are either too badly preserved or too fragmentary

to permit species designation. In fact, 43% of the unidentified specimens are indeterminate

limb shaft fragments generally ranging in size from less than 1 cm to no more than 6 cm,

with the majority less than 3 cm.

Level red deer wild auroch chamois ibex spindet fox Total

E u    
E m      
E l 




Table  : NISP of macrofauna in the Upper Paleolithic and Epipaleolithic levels of Lapa do


The methods used in the analysis are based on those commonly used by

zooarchaeologists. Most of the recovered specimens were piece-plotted while the rest

were recovered in 4mm and 1mm mesh screens. Each piece-plotted bone is kept in its

own plastic bag so the entire assemblage was studied on a piece by piece basis instead of

sorting by part or size category. Identifications were made by comparisons with reference

skeletal material from the Zoology Museum at the University of Wisconsin-Madison, Field

Museum of Natural History in Chicago, and specimens collected in Portugal. Each

identified specimen was recorded according to species or size category, element, portion,

side and age where possible. Limbs were divided into five portions: proximal end, proximal

shaft, middle shaft, distal shaft and distal end. Isolated teeth were recorded separately

from maxillae and mandibulae but all teeth including those still embedded were considered

for estimating MNI. Numbers of identified specimens (NISP) are shown in Table 5.2

In most cases, NISP was the only counting method employed due to small sample

size. The bones from Level F were counted by MNE, MAU and standardized to %MAU.

This level had by far the largest sample size, though it may still be too small for meaningful

statistical analyses. Despite the potential inadequacy, these measures do enable some

important qualitative observations. With regard to limb elements, MNE and MAU was

estimated for each portion. Had epiphyses been the sole portion counted, there would be

almost no limb bone representation. The methods used for counting limb element MNE

was a modified version of that used by Bunn and Kroll (1986) and Marean and Spencer

(1991) (Figure 5.9). Neither estimates of the percentage circumference nor overlapping

tracings of limb shaft fragments were done. The number of identified element fragments

was small enough to visually inspect and compare each piece to determine whether there

was any overlap of specific landmarks such as foramen or muscle attachments or whether

size differences precluded different specimens from originating from the same bone. For

example, as in Todd and Rapson (1988), the proximal humerus shaft is often, though not

always, identified by the presence of the deltoid tuberosity. However, the distal shaft

category would include the proximal-most portion of the olecranon fossa and the

Proximal epiphysis

Proximal shaft


Distal shaft

Distal epiphysis

Figure : Designated limb portions used in the analysis


Figures 5.10: encrusted red deer radii (above) & 5.11: Aurochs astragali (below)

posterolateral nutrient foramen.

Biometric measurements were taken on red deer and aurochs specimens according

to the method used by von den Driesch (1976). These measurements were made with dial

calipers and were recorded in millimeters.

Faunal Remains

Level D

Level D is an Epipaleolithic layer that dates to around 8,300 bp. It was fairly

discontinuous and distinguished from Level E only by its light yellowish brown color.

Fifty large mammal bones were recovered during the excavation. Most were unidentifiable

to species. The few that were include red deer, wild boar and auroch.

Level E

The assemblage from Level E comes from 9 of the 13 excavated units. The level is

characterized by éboulis with whitish gray material that often bonds with the éboulis to

make a breccia. All of the bones from E are covered with a carbonate crust which can,

with care, be removed keeping the surface intact in many cases (Figure 5.10). Fragmentation

is extremely high making species identification more difficult.

In E Upper, dated to 10,070 bp, 183 bones were recovered of which 34 were identified

to species. Twenty belong to red deer with an MNI of only 1, while the other belongs to

wild boar. Both MNIs are based on isolated teeth. Three aurochs specimens were also

identified. Two of these were astragali which were measured to confirm their wild stature.

Both fell well within the wild size range (Bd= 58; GLl= 83.8; Bd= 58.3) (Figure 5.11).

In E Middle, 226 bones were recovered of which 46 were assigned to species.

Twenty-six elements were identified as red deer, with an MNI of 1 and 15 were identified

as wild boar, which are represented by two animals, one adult and one juvenile with an

unfused 2nd phalanx. Five aurochs specimens were also identified.

E Lower contains 107 bones, but this level is much more diverse. Of the 28 elements

identified to species, 19 come from red deer, with an MNI of 1, 7 from wild boar, 1 from

auroch (the sesamoid), and 1 from chamois. The maxillary fragment of chamois contains

the roots of P3, partial P4, and intact M1 (Figure 5.12).

Figure 5.12: Chamois maxillae

Level F

Level F is by far the richest level, with some of the best-preserved specimens and

larger fragments. Much of this level is characterized by loose ebouli with little sediment.

A large hearth found in this level did, however, contain lots of sediment and charcoal

along with many burned and highly fragmentary remains (Figure 5.13). In the 8 units

excavated, 710 large mammal bones were recovered of which 229 were identified to species.

NISP for red deer is 161, followed by wild boar at 67 and one chamois specimen. MNI for

this level is higher, with 4 red deer, 2 wild boar and 1 chamois. The red deer are represented

by 3 adults and 1 juvenile and the wild boar by 1 adult and 1 juvenile (Figure 5.14).

Figure 5.14: Juvenile red deer teeth

Certain parts, the astragali and distal metatarsi , were measured for comparison with

other sites. Metatarsal distal breadth suggests the adult red deer may be represented by a

male and two females. This possibility is discussed further below.


Figure 5.13: bunny pit (above) and 5.15: Level G/I interface (below)

Figures 5.16: refit red deer tibia (above) & 5.17: ibex teeth (below)

Level G

Level G is limited to the area of the large hearth in F. All of the bones are covered

with white, ashy and clayey sediment. Although many are burned or extremely

fragmented, it was rich in bone. Unfortunately, of the 288 recovered, only 47 were

identifiable to species. Twenty-seven of these were red deer and 20 were wild boar, both

with an MNI of 1.

Level I

The stalagmitic crust of Level H has effectively sealed off this level in units D6, D5, E6

and E5 providing the best preservation in the cave (Figure 5.15). There is almost no fine

sediment, only large-sized ebouli. In areas lacking the stalagmitic crust, Level I is in direct

contact with Level F from above. A refit was made of a red deer tibia from I and F (Figure

5.16). This is a reasonable occurrence given the fact that the two levels are distinguished

primarily by ebouli clast size. The large mammal assemblage includes 52 bones. Level I

has represented specimens of 2 red deer and one wild boar. Four ibex teeth and a possible

caprid radius shaft were also recovered (Figure 5.17).

Level J

Level J contains 115 large mammal bones. Most of these come from a small hearth

excavated in 2000 and 2001. The assemblage contains 115 bones of which 26 were assigned

to species. They are predominately red deer with a NISP of 15 and MNI of 1. A few wild

boar specimens and a single auroch phalange were also identified.


Figure 5.18: Photos showing Åmose specimen (above) and Picareiro specimen (below)


The taphonomy of the Picareiro faunal assemblage must be understood before

conclusions can be made regarding human subsistence patterns during the Magdalenian.

Bones deposited in archaeological sites potentially undergo a multitude of destructive

processes from the time of their deposition to the time of the excavation. In this case,

about 12,500 years has elapsed. The primary concerns here are the role carnivores played

in the assemblage formation and whether or not density-mediated attrition due to other

factors has seriously affected the bones. Among these are the chemical weathering of

bones due to exposure, water, root etching and cave geochemistry and the impact of post-

depositional trampling. Once these concerns are addressed, the human behavioral patterns

that led to the deposition of the bones in the first place can be discussed.

The weathering on bones in the assemblage is generally in the form of linear cracks

along the longitudinal axis of long bone fragments, small round pockmarks and eroded

surfaces (Figure 5.18). The weathering cracks point to slow depositional periods in the

cave with bones lying about on the surface for considerable periods of time. Éboulis and

other sedimentation in caves and rockshelters can be notoriously slow (e.g. Laville et al.

1980). The pockmarks seen on many specimens in the éboulis layers are most curious for

they often contain tiny, usually irregular striations running transverse to the grain of the

bone. Nanna Noe-Nygaard (personal communication) has raised the possibility that fungus

and land snails could be responsible for these erosional patterns. Snails have tiny sharp

tongues that could mark bones if they were eating fungus off of them (Miller 1994). The

cave does contain numerous land snails, both in the past and presently, and fungi are

present. The marks present on the bones from Picareiro Cave are very similar to those

documented in the Åmose by Noe-Nygaard (1996) and at Picamoixons in Catalunya by

Verges Bosch (1995). Another possibility is that some kind of acid solution dripping through

from above chemically weathered the bones (Figure 5.19). This is a likely explanation

given that most of the excavated squares are directly below a chimney with no apparent

opening. In addition, the stalagmitic crust forming Stratum H lies directly below the

chimney indicating the flow of carbonate in solution in this area of the cave.

Figure 5.19: chemically eroded wild boar teeth

Carnivore activity

Although Late Pleistocene fauna-bearing sites in Portugal contain carnivore remains,

they are almost entirely absent in the Picareiro assemblage. Only one small tooth, a

premolar of a very young fox was recovered. No postcranial elements are present, even

considering the unidentified specimens, which are all from ungulates of various sizes. A

few bones do show tooth marks, but none that penetrated through cortical bone. The

single auroch specimen in E Lower is acid-etched and could certainly have been passed

by a non-human animal. However, the etching could be natural chemical weathering,

seen on many bones. There appears no doubt that, given the archaeological context of the

bones, that humans deposited most if not the entire assemblage at Picareiro.

The question arises as to whether or not the bones suffered from post-depositional

carnivore ravaging which would obscure human behavioral patterns of butchery and

transport. Bunn (1983, 1986) discussed observations of ungulate body part representation

after hyena ravaging. Hyenas have the ability to crush dense limb bones from most medium

and large ungulates resulting in highly fragmented bone assemblages. Typically, they

completely destroy spongy limb epiphyses such as the proximal humerus, distal radius,

proximal and distal femur proximal tibia and other relatively soft greasy elements (Bunn

1986). The resulting pattern is that midshaft portions are abandoned due to their lack of

grease. Marean and Spencer (1991) and Blumenschine and Marean (1993) conducted

controlled feeding experiments with captive hyenas to determine the patterns of destruction

and remaining element portions in order to understand reverse utility curves seen in many

Paleolithic bone assemblages. These studies are relevant here in spite of the fact that

hyenas went extinct in Portugal by the end of the Last Glacial Maximum because they

were instrumental in understanding why many Paleolithic cave and rockshelter

archaeofaunal assemblages exhibit a reverse utility curve (Figure 5.20) (Marean and Frey

1997; Marean and Kim 1998). Repeated short-term cave and rockshelters occupations by

Paleolithic hunters certainly resulted in the abandonment of large numbers of animal bones

with varying amounts of grease and tissues attractive to hungry carnivores. The two

carnivores known to accumulate ungulate bones are wolf and fox, both of which were

present in Portugal during the Late Pleistocene. While the former is easily capable of

hunting large game, the latter cannot bring down adult medium-sized ungulates. On the

other hand, domestic dogs, present already by the Late Pleistocene in other parts of Europe,

may have been the secondary consumers of these bones. Therefore, it is important to

consider actualistic and experimental studies of modern wolves and dogs to understand

the patterns of bone damage and destruction these animals create.

Bulk Strategy
Gourmet Strategy



Gourmet Strategy
Food Utility Index Food Utility Index

Figure  : Schematic illustration of utility curves (A) represents the skeletal parts removed from
kill/butchery sites (B) represents parts left at kill/butchery sites (after Metcalfe and Jones  )

Payne and Munson (1982) cite the work on Mesolithic sites by Steenstrup (1862) as an

example of the early recognition that dogs in prehistoric Europe affected skeletal part

representation by consuming the weaker portions of bones. Ethnographic observations

of Eskimo dog feeding confirmed Stenstrup’s suspicions (Steenstrup 1862: cited in Payne

& Munson 1982). The most well known ethnoarchaeological studies of bone destruction

by canids are those by Brain (1967, 1981), Binford (1981), Walters (1984) and Hudson (1993).

Brain (1967, 1981) observed damage to and survivorship of goat bones discarded by

Hottentot villagers in southern Africa which were then further damaged by their dogs.

Binford (1981) has provided details on damage to caribou and sheep bones by wolves.

His data show that as the degree of fragmentation increases (measured in #s of splinters/

MNI), the number of articular ends missing increases (Binford 1981: Figure 4.59). The

only case where Nunamuit created a similar pattern was after bone grease processing.

Payne and Munson (1982) conducted controlled feeding experiments with large dogs

during which they fed a hungry Ruby and her son, Slick, squirrels, rabbits and parts of

goats. The pattern of destruction on the goat bones are of interest because of their

comparable size to the animals represented at Picareiro. The dogs were fed half a goat on

one occasion and limbs-only on a second. The pattern of destruction was a reduction of

most parts to small ~2cm-sized fragments. The survivorship of limb portions was

comparable to that observed by Brain among the Hottentots. Survival was low for the

scapula, proximal humerus, proximal and distal femur, proximal tibia and high for the

distal humerus, proximal radius, distal tibia, proximal and distal metapodials. Two juvenile

goat heads were fed to the dogs and both the maxillae and mandibles were destroyed

leaving only half of the deciduous teeth. Walters (1984) provides additional

ethnoarchaeological data on bone attrition by dogs at hunter-gatherer sites in Australia.

However, he did not discuss the resulting skeletal element pattern. Both Walters and

Hudson (1993) noted the overwhelming destruction and loss of bones from small game

animals by dogs in ethnoarchaeological bone assemblages. The abundance of rabbit bones

in the Late Upper Paleolithic caves of Iberia suggests dogs or other hungry carnivores did

not impact small animal bone assemblages.

During the Paleolithic, wolves and dogs likely gained access to ungulate skeletal parts

stripped of meat and cracked open for marrow. Greasy limb epiphyses may have been

discarded or pounded, ground and/or boiled to extract additional fat. Depending on

circumstances some elements may have been preferred over others leaving only the most

marginal parts to the dogs. The expectations for skeletal part representation if dogs affected

the bone assemblage are that the less dense limb epiphyses would be absent (Figures

5.21a&b), ribs and vertebrae would be damaged, the pelvis gnawed to the acetabulum,

skull parts including the mandible would be damaged or destroyed, metapodials would

exhibit moderate damage, phalanges would be complete, carpals and tarsals would exhibit

acid-etching, there would be a high degree of bone chips with pitting and polishing and

limb shafts may survive in greater abundance.

Bone density

Given the lack of carnivore remains and possibility that companion dogs ravaged the

bones in Picareiro, it is important here to evaluate the likelihood that density-mediated

attrition due to other factors affected the Picareiro faunal assemblages. Bone attrition,

whether by carnivores or other natural factors, is highest in the portions with the highest

fatty acid content (Lam et al. 1999). This makes them attractive to carnivores for their

nutritional value but structurally weak. Even in the absence of carnivores, their destruction

Figures 5.21a &b: Carnivore-gnawed ungluate limb bones from Portugal


is still more likely. A great deal of work has been done since Brain’s (1967) pioneering

comparisons of bone density to skeletal part frequencies in human and carnivore-

accumulated faunal assemblages. Lyman (1984) used photon densitometry to measure

bone mineral densities of deer, pronghorn and sheep bones in order to understand the

affect bone density has on skeletal element patterning in archaeological contexts. Since

then many other animals have been measured using photon densitometry (PD) including

bison (Kreutzer 1992), llama, guanaco, vicuña (Elkin and Zanchetta 1991; cited in Lyman

1994), seal (Chambers 1992; cited in Lyman 1994) and small animals such as marmot (Lyman

et al. 1992) and rabbit (Pavao and Stahl 1999). Though variability in bone volume density

occurs among ungulate species, there are regular patterns in the volume density (VD) at

the scan sites. Mandibles are relatively dense with the exception of the ascending ramus.

In deer, pronghorn and sheep, vertebrae are generally weak. This contrasts with bison,

which has much higher VD measures, likely due to methodological differences in the

calculation of volume density by Lyman and Kreutzer (Lam et al. 1999). The same could

be said for ribs and parts of the pelvis. In the forelimb, the scapula, proximal humerus,

ulna and phalanges have a relatively low density. The radius, carpals and metacarpals

have some of the highest density values. In the hindlimb, the proximal and distal femur,

proximal tibia and proximal calcaneus are relatively low density. The rest of the tibia

(especially the midshaft distal to the crest), tarsal, astragalus, calcaneus and metatarsal

(with the possible exception of the distal shaft) have the densest parts. Recent work by

Lam et al. (1998) using computed tomography (CT) scans of bones from horse, wildebeest

and reindeer shows some important differences between the two methods. However,

they also observed a strong degree of similarity in bone mineral density values between

artiodactyls and perissodactyls. Therefore, any differences in skeletal element

representation among species in archaeological sites is likely due to other destructive agents

such as carnivore ravaging or human butchery and transport patterns. Although no studies

have been done on Cervus the density values for other artiodactyls are considered

representative here. Ioannidou (2003) has just published results of X-ray densitometry

scans of adult and juvenile wild boar and domestic pig. Those values are used here to

evaluate wild boar survivorship. Her comparisons with sheep and cattle revealed

significant inter-taxonomic differences in bone density. Bone density differences were

also found between young and adult wild boar, adult males and females and wild boar

and domestic pig.

Within the limb elements some further observations are apparent from both PD and

CT scans. In CT scans 6 out of the top seven in the rank of long bone scan sites are midshafts

(Lam et al. 1998: Table 2). Compared to PD, the midshaft rank for radius, humerus and

femur is considerably lower. Where femur ranks highest in the CT scan method, it only

ranks 17th in the PD method. The humerus ranks 4th in the CT method but only 11th using

photon densitometry. The differences in scan site rank are probably due to the way in

which each method treats cortical bone thickness (Lam et al. 1999). In general, both methods

demonstrate that midshafts are the densest portion of limb bones followed by proximal/

distal shafts and epiphyses. Therefore midshafts are the most likely portion of a limb

element to survive density-mediated destruction.

The importance of counting midshafts in the calculation of Minimum Number of


Elements (MNE) has been debated in the literature for two decades (Bunn 1983, 1986,

1991; Bunn & Kroll 1986; Bunn et al. 1991; Binford 1986; Marean & Spencer 1991; Stiner

1994, 1998, 2002; Marean & Frey 1997; Marean & Kim 1998; Bartram and Marean 1999;

Klein 1999; Marean et al. 2001). Hill (2001) used the volume density estimates derived

from CT scans to develop three expectations regarding limb bone representation and

density-mediated attrition. These are:

1) a midshaft landmark will yield the highest landmark MNE

2) upper long bones will outnumber metapodials, and

3) the femur will be the most abundant long bone (Hill 2001: 121).

In order to test whether the bones from Picareiro have suffered density mediated attrition,

the %survivorship or %MAU was plotted against bone mineral density values from Lam

et al. (1999) (Figure 5.22, Figure 5.23). The %MAU was calculated by taking the MNE and

dividing by the number of occurrences in the skeleton to derive the MAU. The MAU was

standardized by converting the highest MAU to 100 and dividing each MAU value by the

value of the highest one (Binford 1984).

For Picareiro Level F, the red deer %MAU are plotted against the BMD of caribou

from Lam et al. (1999). Figures 5.22 and 5.23 shows the scatterplots and values for Pearson’s

r and Spearman’s rank correlation coefficients. The values are roughly equivalent (r=

0.47, p; rs= 0.50, p<.001) and show a weak but significant positive correlation between

bone survivorship and bone mineral density at Picareiro. Density-mediated attrition or

carnivore-ravaging may be responsible for some loss but neither is the primary factor in

LP Level F red deer







.2 .3 .4 .5 .6 .7 .8 .9 1 1.1 1.2

Figure 5.22: Plot of skeletal element frequency vs bone mineral density (CT)

LP Level F red deer







.1 .2 .3 .4 .5 .6 .7 .8
bone density (PD)

Figure 5.23: Skeletal element frequency vs bone density (PD)


skeletal element representation at Picareiro. Hill’s (2001) expectations for bone density

are met by the first two (highest MNE for long bones is based on midshafts and upper

limb bones outnumber metapodials), but not the third since the femur is not the most

abundant long bone.

Skeletal element representation and butchery patterns

One of the most debated topics in zooarchaeology centers on the bones left behind by

prehistoric hunters at kill sites and carried away and discarded at base camps. Bunn

(1991) traces the origins of this problem to the work of the early prehistorians Lartet and

Christy (1865-1875) who first suggested that the skeletal part frequencies of the French

Paleolithic sites reflected the transport decisions of prehistoric hunters. They argued that

for larger animals, the axial elements were discarded at kill sites while meat and marrow-

rich limb elements were transported to residential camps. Medium-sized ungulates such

as reindeer were transported whole. This latter observation is more relevant to the Picareiro

assemblage and is discussed further below. In Americanist zooarchaeology the issue was

renewed by White (1953) who considered anatomical part frequency in determining

whether a site represented a kill or a residential site. Perkins and Daly (1968) argued that

an overrepresentation of foot bones in a site occurred because hunters stripped the meat

off the limbs, presumably to reduce the weight, and carried it back to the residence in the

skin of the animal with the feet still attached for use as handles. The bulky limb bones

were left behind. In the decades since, zooarchaeologists have realized there is a great

deal of variability in bone assemblages and numerous causes for their composition. There

is no longer a simple dichotomy between kill sites and residential sites (Binford 1978,

1980, 1981). Ethnoarchaeological research has shown many factors determine which parts

of the skeleton are transported from kills and which ultimately arrive in camps and/or

villages (Binford 1978; Bunn 1986, 1991, 2002; Bunn et al. 1988, 1991; O’Connell et al. 1988,

1990; O’Connell and Marshall 1989; Bartram 1993; Emerson 1993; Oliver 1993). What

happens along the way and in camp is discussed in the next section.

Binford (1978) quantitatively modeled transport decisions by devising body part utility

indices for caribou and sheep. These indices, Meat Index, Marrow Index, Grease Index,

were combined to produce a general utility index (GUI) that was modified (MGUI) to take

into account the low-utility parts attached to high utility ones whose removal coasts

outweighed the benefit (reducing transport costs) of removing them. Generally, high-

utility skeletal parts will be missing from kill sites because hunters would have processed

the carcass and left behind low-utility parts. Each of the indices has been further modified

by others but serves as the basis for many behavioral interpretations of archaeofaunal

assemblages. This model of discarding low-utility axial parts and transport of high-utility

appendicular parts has been challenged recently by O’Connell and colleagues who used

ethnographic observations of Hadza hunters to argue that for many large game animals,

appendicular parts are left behind more often than axial parts, which are transported away

(O’Connell et al. 1988, 1990; Emerson 1993). The reasoning behind their argument is that

limb bones are heavy and hunters will strip the meat off of them and discard the bones,

taking the axial parts back to camp because the effort to remove the meat from them

would be uneconomical. The decision to transport a skeletal part is guided by the ratio of

edible to inedible tissue. Bones with a low ratio are likely to be processed to remove meat

and marrow and the bulky, heavy inedible fraction, bone, will be discarded. Elements

with high ratios would be transported because the cost to remove the inedible fraction

would lower the economic value of the edible part. O’Connell et al. (1988) argue transport

decisions are based on carcass size and distance to camp. Not only do they question the

White model and claim the reverse (Perkins and Daly), they also call into question the use

of body size class as an analytical unit, pointing to their data that suggest variation in the

treatment of carcasses within size class (O’Connell et al. 1990). Bunn (1993) countered this

interpretation of Hadza transport by arguing that Hadza hunters try to transport entire

carcasses when possible. Exceptions are very large animals (e.g. giraffe), those that have

poor marrow quality and decisions to avoid sharing high-utility parts such as rich, marrow-

filled limb bones (Bunn 1993). The high proportion of vertebrae is due to the ability to boil

and render grease from them. Thus they may have a higher economic value and greater

incentive for transport than they may have in pre-boiling times such as the Plio-Pleistocene.

This technology was probably available to Upper Paleolithic hunter-gatherers and is a

relevant factor to interpret assemblage composition at Picareiro.

Bartram (1993) reported ethnoarchaeological observations of carcass transport by Kua

hunter-gatherers. His study showed an important correlation between assemblage

composition at kill/butchery sites and time spent field processing. Large game limb

elements were left at kill/butchery sites after meat was stripped off and sun-dried. This

occurred because of decisions to reduce transport costs based on factors such as the number

of carriers, distance to camp and size of the animal. Stripping and drying meat was one

possibility among several.

The debate over reducing transport costs usually centers around animals of large to

very large size (>200kg). In Portugal, large game available during the Late Pleistocene

include horse (700-800 kg), aurochs (500-800 kg?), red deer (100-180 kg), roe deer (15-30

kg), ibex (35-80 kg), chamois (30-50 kg) and wild boar (80-100 kg) (not shown: Table 5.3).

The first two fall into the size group 4 of the large African bovids and equids hunted by

the Hadza and Kua. The other five are smaller and fall into the range of size group 2-3

animals typically transported as entire cut up carcasses.

The statistical correlations between carcass size, distance to kill and proportion of

elements transported were argued by O’Connell et al. (1990) to explain 40-57% of the

variation. Although the correlations were significant the fact remains that they only explain

half of the variation. The remaining variation is significant and just as important in

determining carcass transport. The additional problems of sample size and length of

observation time was raised by Bunn (1993) who provided data from 110 carcasses. The

transport pattern shows that over 90% of the carcass was transported with the exception

of heads and ribs which were carried over 80% of the time. Foraging models that attempt

to simplify decision-making to one or two variables, such as the amount of edible tissue,

the amount of processing time, distance traveled and energy expenditures always make

the qualifying statement “all else equal.” Rarely, does it seem, that all else is equal regarding

human decision-making.

The skeletal element representation for Level F red deer and wild boar at Picareiro

Level F

Wild Boar NISP MNE

Table : Picareiro Level F wild boar
(isolated teeth not included)

Figures 5.24a &5.24b: Wild boar specimens from Picareiro


shows some interesting patterns. Table 5.4 and 5.5 shows NISP and MNE for wild boar

and red deer, respectively. It is immediately apparent that red deer has a much better

overall skeletal representation than wild boar. While Level F has evidence for at least 2

wild boar and maybe 3, the NISP and MNE show an extremely high degree of under-

representation for most elements.

Rowley-Conwy et al. (2002) have recently published a food utility index (FUI) for

European wild boar. Using the revised methodology proposed by Metcalfe and Jones

(1988), they found the highest utility parts were the entire thorax (vertebrae, ribs and

sternum combined), lumbar vertebrae, pelvis and femur. Of secondary importance was

the skull, mandible, cervical vertebrae and scapula. The humerus, tibia and lower limb

elements ranked lowest. This contrasts with the values Binford (1978) reported for sheep

and caribou. Rowley-Conwy et al. (2002) show that wild boar femora, scapulae, humeri

and tibiae generally have a much lower food utility than caribou especially. In Picareiro

Level F, crania and mandibles are the most common parts, followed by phalanges and

metapodials (Figures 5.24 & 5.25). The vertebrae, ribs and upper limbs are least represented.

The tibia, humerus and ulna are the only upper limbs present. This suggests removal of

the bulky heads and transport of nearly intact carcasses away from the cave.

Some wild boar was consumed onsite as evidenced by the few limb remains. It is

unlikely that meat was removed in order to reduce transport costs. Drying and smoking

of limb quarters may have taken place, but likely with the bones in. Based on the wild

boar FUI, it would appear that the Picareiro wild boar exhibit a classic reverse utility

curve for kill/butchery sites where the lowest utility parts are most commonly left behind.

Table : NISP of Red Deer from Magdalenian levels in Lapa do Picareiro

Red Deer Level E u Level E m Level E l Level F Level FA Level G




P Hu epi      
P Hu shaft      
Hu midshaft      
D Hu shaft      
D Hu epi      

P Ra epi      
P Ra shaft    
Ra midshaft      
D Ra shaft      
D Ra epi     

P Ulna      
D Ulna      


P mtc     
Mtc midshaft     
D Mtc     


P Fe epi      
P Fe shaft      
Fe midshaft      
D Fe shaft    o  
D Fe epi      


P Ti epi      
P Ti shaft     
Ti midshaft     
D Ti shaft      
D Ti epi      


P Mtt     
Mtt midshaft   
D Mtt      


Table : Skeletal element representation for red deer Level F

Side and age taken into consideration in MNE estimation
Level F



V cervical   
V thoracic   
V lumbar  

Prox Hum epi    
Prox Hum shaft  
Hum midshaft    
Dist Hum shaft   
Dist Hum epi    
Prox Rad epi    
Prox Rad shaft   
Rad midshaft   
Dist Rad shaft   
Dist Rad epi  
Prox Ulna    
Dist Ulna    

Prox metacarpal    
Metacarpal midshaft   
Dist Metacarpal  

Prox Fem epi    
Prox Feh shaft    
Fem midshaft    
Dist Fem shaft    
Dist Fem epi    

Prox Tibia    
Prox Tibia shaft   
Tibia midshaft   
Dist Tibia shaft   
Dist Tibia epi    

Prox Metatarsal    
Metatarsal midshaft   



: Picareiro Level F red deer complete bone element
repres entation

Level F



V cervical  

V thoracic   
V lumbar  






Table  : Picareiro Level F red deer skeletal element representation

(limb shafts not counted)
Level F



V cerv   
V thor  

V lum  

P Humerus   
D Humerus   
P Radius   
D Radius 


P Metacarpal    
D Metacarpal  
P Femur   
D Femur   
P Tibia    
D Tibia   

P Metatarsal    
D Metatarsal  


Table 5.6 shows MNE, MAU and %MAU for red deer. The crania are predominately

represented by teeth with a few intact maxillae and broken mandibles. Table 5.7 shows

MNE estimates for five limb portions contrasted with MNE estimates for limbs using

epiphyses only. Clearly, element representation would be much different had limb shafts

not been taken into consideration. In fact, counting MNE by epiphyses-only leads to a

classic reverse utility curve for kill sites (Figure 5.20) (Binford 1978; Metcalfe and Jones

1988). This could lead to an erroneous conclusion that Picareiro itself was a kill site,

dominated by low-utility skeletal parts with most of the limbs transported away, or that

carnivores had ravaged the assemblage. Meaty limb elements are in fact present and

mostly identified by shaft fragments with few surviving epiphyses. Only dense epiphyses

of lower limbs are well represented. The missing epiphyses are the proximal humerus,

proximal radius and ulna, proximal and distal femur and proximal tibia. These are precisely

the low density but greasy parts most likely to disappear due to natural attrition or carnivore

consumption. With limb shafts counted the overall representation is shown in Table 5.7.

%MAU shows a high percentage of upper limbs except the ulna and femur. Lower limbs

are moderately represented by comparison with the exception of smaller dense carpals

and tarsals. However, in the indeterminate species fraction there are a number of carpals

and tarsals not identified to species but assignable to a Cervus-sized or medium-ungulate

size class. These were smaller than the reference specimen (female Cervus canadensis) and

probably came from smaller red deer. In any case, the most abundant elements are limbs

while ribs, vertebrae and pelves are scanty. Even counting fragments from the

indeterminate species fraction, ribs and vertebrae are very much underrepresented. This

could be due to some density-mediated attrition or carnivore-ravaging, though none of

these show evidence for carnivore gnawing.

Figure 5.27: red deer femora fragments

As with the indeterminate fraction, the identifiable assemblage in each Magdalenian

level at Picareiro (and even the undated lower levels) shows a high degree of fragmentation.

Patterns of ungulate long bone fragmentation have been used interpret faunal assemblages

and infer human behavior for several decades (Binford 1978, 1981; Brain 1981; Bunn 1983,

1986; Speth 1983; Bunn and Kroll 1986; Todd and Rapson 1988). In Picareiro, the long

bones show evidence of intentional cracking for marrow extraction. They are highly

fragmented with many percussion scars and impact fractures (Figures 5.27; 5.28; 5.29).

None of the long bones are complete. The only whole elements are 3rd phalanges,

Figures 5.28: red deer humeri fragments & 5.29: red deer lower limb fragments

carpals, tarsals, calcanei, astragali and sesamoids. Every limb element including 1st and

2nd phalanges of red deer are cracked open for marrow extraction. The frequency of small

(<2cm) fragments suggests dry bone breakage due to post-depositional trampling of

sediment, mostly éboulis. Cutmarks are difficult to assess due to the chemical erosion on

many of the bone surfaces.

Figure 5.30 shows complete bone %MAU vs (S)FUI for Level F red deer (Figure 5.31a

shows the relationship when limb shafts are not counted). Although the correlation is

negative it is insignificant. Grayson (1988) has argued that positive significant correlations

for %MAU and bone density coupled with insignificant correlations between %MAU and

utility indices means that density-mediated destruction and not differential transport is

responsible for the assemblage composition. Using Lyman’s (1994: 264) chart for possible

combinations of correlation coefficients, it would appear that the Level F red deer fall into

his class 4, a ravaged assemblage. As Lyman points out, however,

indices of utility, transport and structural density of bone parts are ‘frames
of reference,’ and as such their use as explanatory algorithms for skeletal part
profiles is not advised; rather, they should be used as one of several steps in
building a taphonomic explanation for those profiles (Lyman 1994: 281,
emphasis in original).

Given the absence of carnivore remains, paucity of carnivore gnaw marks and low

impact of density-mediated attrition, absence of limb epiphyses and vertebrae suggests

grease extraction. In fact, Binford (1981) discussed the relationship between long bone

splinters and articular ends. In carnivore-ravaged assemblages, splinters represent “what

is no longer present,” which is the articular ends (p.177). In assemblages left by humans,

splinters represent “what remains,” which again are the articular ends. The exception for

LP Level F red deer







0 20 40 60 80 100

Figure 5.30: Skeletal element frequency vs standardized

complete bone food utility

LP Level F red deer







0 1000 2000 3000 4000 5000

Figure 5.31a: Skeletal element frequency vs unstandardized food utility

Proximal and distal ends using limb epiphyses only

humans is grease extraction whereby articular ends are destroyed thus mimicking a

carnivore-ravaged assemblage. Preservation bias due to density-mediated attrition may

have impacted ribs and vertebrae since they are present in very low proportion. However,

given the above discussion of carnivore destruction, grease processing and

ethnographically observed carcass transport these elements may be absent for other reasons.

Both Bunn (1993) and Bartram (1993) observed the removal and consumption of ribs by

Hadza and Kua hunters at kill/butchery sites.

Transport of elements is difficult to assess given the extremely fragmentary nature of

the bones. However, results indicate an equal representation of limbs with the possible

exception of femora. It is possible that meat was filleted, dried or smoked in the cave and

transported away. However, since cutmarks are rare this is difficult to test. The remaining

bones were then cracked open and the marrow consumed onsite. This would appear to

contrast with the Hadza model of carcass treatment if Picareiro is considered a butchery

site. Obviously it is not a kill site but a place to which animals were transported and

processed. Given the size class of the animals hunted, all elements should be represented

if entire carcasses were brought to the cave and consumed onsite. The likelihood that the

axial portions of animals of such size would have been left at kill sites is low (Bunn 1991,

1993). However, Bartram’s study of Kua carcass treatment provides some interesting

possibilities. The number of skeletal elements left behind at Kua kill and/or butchery

sites correlated positively with time spent processing. If Magdalenian hunters spent a fair

amount of time in Picareiro during each visit that may account for the degree of carcass

processing of medium-sized ungulates. Increased occupation time may have led to filleting

and drying/smoking of meat from limb elements. Epiphyses and vertebrae may have

been carried back to a base camp for grease extraction. The absence of femora and pelves

provides further evidence of transport. Binford (1978) noted low frequencies of femora at

butchery sites or field camps. A similar pattern was observed by Noe-Nygaard (1977) for

Star Carr, though Binford (1981) hypothesized that it was due to site function. David and

Farizy (1994) show an under-representation of femora and pelves at Mauran, a Middle

Paleolithic bison kill site in France. This site shows a strong positive correlation between

%MAU and bone density (PD) and a reverse utility curve for %MAU vs MGUI and %MAU

vs RGI (Revised Grease Index) (David and Farizy 1994). Given that limb shafts were

counted in MNE estimation Mauran could represent a site with density-mediated attrition

or a kill/butchery site where high utility parts were transported away or destroyed during

grease production.

Figure 5.33: red deer 1st phalange with impact fracture


In addition to limb bones, the 1st and 2nd phalanges also show evidence for marrow

extraction (Figures 5.33, 5.34 & 5.35). They are all split open and exhibit impact fracturing,

similar to ones documented at Erralla (Altuna and Mariezkurrena 1985) and La Riera

(Altuna 1986) in Cantabria, at Tossal de la Roca (Perez Ripoll 1992) and Cova de les Cendres

(Villaverde and Martinez Valle 1995) in Valencia and at Cingle Vermell (Vila et al. 1985) in


Binford (1978) suggested this was a sign of subsistence stress due to the effort

required to obtain a small amount of marrow. He observed Nunamiut processing of caribou

phalanges only during periods of food shortage and by “old timers” (Binford 1978). Jones

and Metcalfe (1988) simplified Binford’s Marrow Index and ranked return rates of caribou

body parts. They concluded that despite the high fat content (% oleic acid), phalanges

offered the lowest return rates (calories/hour) due to their low cavity volume, caloric

content and relatively high processing times. Their expectations derived from optimal

foraging theory seem to match the marrow processing patterns Binford observed among

Nunamiut hunters. However, the scapula and mandible were also ranked low and not

processed by the Nunamiut. The mandible was commonly broken open for marrow

extraction by Upper Paleolithic hunters across Eurasia. Jones and Metcalfe (1988) calculated

the caloric content of marrow using a constant value of 4 kcal/ml. Since the stated range

(0-9 kcal/ml) reflects lipid content fluctuations in bone marrow, the return rates for each

element will fluctuate as well. This will not likely affect the ranking of parts but it will

increase the value of “low-ranked” parts such as mandibles and phalanges during certain

seasons. Of course, the counter argument would be that low ranked resources will not be

Figure 5.34: red deer 1st phalanges split to extract marrow (above) &
5.35: 2nd phalanges

incorporated into the diet unless high ranked ones are no longer available or at least reduced

to a certain point. Therefore, according to the diet breadth model, phalanges and mandibles

will always be ignored unless overall availability is low. Does the cracking of mandibles

and phalanges represent resource depression? Or, does the use of rifles and snowmobiles

to hunt caribou inflate the ability of Nunamiut hunters to procure animals to the point

that they no longer need to process items that were in the past when technology constrained

resource yields? It is not surprising that “old-timers” would still process these parts if

they had done so regularly in the past. Still, it could mean that their access to carcass parts

was restricted by younger hunters and they were forced to process parts that otherwise

may have been ignored or commonly utilized by non-hunting members of society (e.g.

women, children and elderly). Jones and Metcalfe (1988) conclude that mandibles, pelvis,

scapula and phalanges will be processed in rank order as the nutritional quality of an

animal decreases. Therefore, one would expect greater use of these items during stressful

periods like the late Winter and especially the early Spring when the fat reserves of

ungulates are depleted. Jones and Metcalfe (1988) argue that phalanges would be the last

element to suffer fat depletion. Thus, as the amount of fat decreases in each limb element

the more valuable phalanges become. Considering that Fall and Winter are not usually

thought to be times of dietary stress since animals would have been at their fattest during

the Fall through early Winter, the processing of phalanges (and mandibles) for marrow at

Picareiro would appear to reflect other factors (Speth & Spielmann 1983). In the

Mediterranean region, summer is likely to be the most stressful season given the rainfall


One possibility could be related to sexual dimorphism. Hunters targeting males

could resort to extracting marrow from the phalanges if they were killed during a season

of pronounced fat depletion. Food utility indices for sexually dimorphic species should

be modified to account for differences in fat content between males and females (Speth

1983; Metcalfe and Jones 1988). Most ungulate females retain higher amounts of fat reserves

during winter because of reproductive needs. Therefore, late Winter and early Spring

females may also have higher marrow and grease values in limbs compared to males.

Perhaps instead this is a signal for large game resource depression. Proponents of

optimal foraging theory would argue that the intensive ungulate carcass utilization is a

sign of resource depression (e.g., Broughton and O’Connell 1999). Furthermore, they would

argue that the high number of rabbit carcasses deposited in caves and rockshelters during

the Magdalenian adds further support for resource depression because small game ranks

low and would only be included in the diet if higher ranked resources were not available

in sufficient quantity.

Comparisons between the treatment of large and small game lead to further

hypotheses. Hundreds of rabbit carcasses were intensively processed at Picareiro (Hockett

and Bicho 2000) (Table 5.9a & b). Carcasses were roasted whole since only the feet have

high frequencies of burning. Though cutmark frequencies were not analyzed, the meat

was probably removed from the limbs before they were broken open to extract marrow.

Almost certainly rabbits were consumed onsite in their entirety. The low frequency of

vertebrae suggests they were probably ground into a fat rich meal, a practice common in

the Great Basin of the United States (Hockett 1991). Whether this was done onsite or at

Table a: Number of Rabbit Bones per Stratum

at Lapa do Picareiro



E (UPPER)   







Table b: rabbit elements at Picareiro

A D E F G/I J Total
Upper Lower












base camps is unknown. Although the rabbits represented at Picareiro were consumed

locally, we do not know how many may have been procured and transported away from

the site. The evidence of repeated use of the large hearth or “bunny pit” suggests the

bones do not represent the number of animals processed in the cave. Level G is essentially

a finely laminated ash deposit showing multiple uses over a long period. The overlying

Level F hearth may only represent the final one or superimposed few uses before disuse.

Therefore, it is likely that many more rabbits were procured near the site.

Why were hunters at Picareiro? Was it a specialized hunting camp for procuring

red deer, wild boar and rabbit? Were the primary goals to maximize meat yield or large

game fat and protein yield? Was it to collect rabbit and opportunistically hunt red deer,

caprids and wild boar? Or, hunt large game and trap rabbits for immediate consumption?

Everything brought in seems to have been consumed. Did ungulate resource depression

lead Magdalenian hunters to intensively harvest and process large numbers of rabbit?

This question requires knowledge of the seasonal use of the cave and comparisons of

large game and rabbit carcass utilization from additional sites in the region.


Preliminary seasonality determinations were made using cementum annuli analysis

on red deer teeth (Klevezal 1996; Pike-Tay 1991). Work was carried out with Tina Dudley

in the MacDonald Institute for Archaeological Research at Cambridge University. Modern

specimens from the Scottish Highlands were used as a control sample. Two maxillary M1

teeth from different animals in Stratum F showed late Fall/ early Winter season of death.

Figure 5.36: Wild boar phalanges from Picareiro Level F

Figure 5.37: distal wild boar tibia with fusion line still visible

Additional determinations were attempted based on the timing of 2nd phalanx

epiphyseal fusion in modern wild boar from research by Bull & Payne (1983) and Bridault

et al. 2000) (Figure 5.36). This is much more tenuous given the wide range of time estimates

between those authors and vagaries in the definition of unfused/fusing/fused. The

specimen from Level F is clearly unfused, and would fall within the Bull and Payne 7-11

month category while the specimen from E Middle with a fusing epiphysis, would fall

within the 19-23 month category. The Bridault et al. (2000) data from 48 wild boar killed in

France have filled the gaps in the Bull and Payne data. The second phalanx is unfused

until about 9 months when fusion begins. Complete fusion (line invisible) is achieved by

18 months. Therefore, the unfused specimen in Level F would be less than 9 months old.

Had the birth taken place in February, the animal was killed sometime before October.

An April birth would but the death sometime before January. Season of death can not be

determined reliably because the animal could have been killed any time during the year.

The fusing specimen in E Middle came from an animal 9-17 months old. Thus, if the

animal were born in February, it died sometime between the following October and July.

An April birth would put the season of death between the following January and

September. The wide range in fusion time for the second phalanx makes this element an

unreliable seasonality indicator.

A distal tibia specimen has a fused epiphyses with fusion line still visible (Figure

5.37). This falls within the Bridault et al. (2000) 17-25 month range so that an animal born

in February may have been killed between July and March of the following year. An

August-born one would indicate a death between January and September. While

epiphyseal fusion rates provide useful and reliable age at death estimates for the young

wild boar the potential of females giving birth a second time during the year precludes

their use for determining season of death. More research using stable isotopes may help

determine whether wild boar had two births in the Late Pleistocene (e.g. Balasse et al.

2003). Currently, a more reliable indicator may be the population dynamics observed in

modern wild boar. The occurrence of both mature adult males and juveniles would suggest

the animals were killed during the reproductive season when adult males join with females.

This would imply a late Fall/Winter/possibly early Spring season of death for wild boar

at Picareiro.


The other archaeological levels at Picareiro do not have comparable macrofaunal

assemblages to Level F. This is probably due to the lack of substantial features like the

large hearth or bunny pit. Level J contains a small hearth but it is not yet absolutely dated

and there are no fossile directeurs to assign a technological phase (Figure 5.38). Aurochs is

present in J though their remains are limited to foot bones. What happened to the rest of

their carcasses? There is no indication in the unidentified fraction that any aurochs limbs

were left in the cave. The presence of auroch is strange given the elevation of the cave (540

asl). These animals are thought to be lowland plains or forest-living animals, therefore

the represented parts should not indicate an auroch was killed nearby and processed so

that only feet were brought into the cave. The elements came directly out of the small

hearth so it would seem that humans were responsible. Is it possible that a carnivore

brought them in and left them inside an abandoned hearth? Perhaps they were bones

associated with a foreleg snack brought by one of the hunters.

Figure 5.38: Level J hearth in front of south profile

The four ibex teeth recovered in Level I probably indicate a cooler climatic phase

but the lack of absolute dates and diagnostic lithics precludes the assignment of this

occupation to a cultural period. No substantial features like the Level F hearth were

found in this level. The occupation during this period was very short.

The Magdalenian faunal assemblages from Levels E, F and G are numerically

dominated by rabbit. Large numbers of rabbit carcasses were processed and consumed

onsite. Entire carcasses of medium ungulates were brought to the cave for processing.

Many red deer limb elements were stripped of flesh and the bones broken open to extract

marrow. Epiphyses were either consumed by dogs or pounded and boiled (?) to extract

grease onsite or at a base camp. Wild boar was probably brought to the cave where heads

and some lower limb elements were consumed. The rest of the carcass was probably

transported away. Therefore, it appears that wild boar were preferentially transported

over red deer. This may have been due to nutritional differences between the two. Table

4.12 showed wild boar consistently has more fat than red deer. This may have been the

primary motivation for transporting most of the wild boar meat and marrow back to

residential camps. The same might be said for chamois but the sample is extremely small.

The presence of aurochs is perplexing since the cave is not located near preferred habitat.

Marine fish, sardine and shad, are also present in the Magdalenian levels (Bicho et

al. 2000). These may have been smoked or dried and carried in as hunters’ snack food.

Belcher’s analysis (Bicho et al. 2000) shows a pattern of small fish consumption analogous

to that observed ethnoarchaeologically where heads are chewed up and the vertebrae

removed and discarded. A small number of marine clam, scallop, limpet, mussel and

gastropod shells are further indication that the coast was within the economic catchment

of Picareiro. Their presence is not surprising given the fact that other sites nearby contain

small shellmiddens dated to the Early Holocene. These are discussed later.

In sum, the extraordinarily high number of rabbit bones in the large hearth, carcass

butchery and high degree of fragmentation of the large mammal limb elements all indicate

the cave was repeatedly used as a hunting/carcass processing camp by groups of hunters

from larger residential sites in the surrounding valleys. Seasonality information suggests

visits generally occurred during the late Fall and early Winter. The data are too limited to

know if the appearance of fish and additional ungulates in the upper levels represent

overall dietary diversification. This is almost certainly an artifact of sample size. The

Picareiro data alone are not sufficient to test the Broad Spectrum Revolution model or the

Bicho and Zilhão models. These require additional information from other sites in the


5.4 Lapa do Suão

Lapa do Suão is located in a narrow limestone valley about 40 km north of Lisbon

and approximately 15 km from the Atlantic Ocean. The cave lies about 50 m above the

Roto valley floor and its opening faces NE (Zilhão 1997) (Figure 5.40). The cavity has a

long passage with a SW inclination. There is a 25m2 room at the end.

The cave was first investigated in the late 19th century by the geologist Carlos Ribeiro

(Rocha 1907). Santos Rocha viewed the materials collected by Ribeiro in the cave entrance

and concluded that they were from the Iron Age (Rocha 1907). In the 1960s, Furtado et al.

(1969) excavated Neolithic and Calcolithic deposits in the entrance and passage. They

reported finding numerous microliths, possibly Mesolithic, and a pointe à cran or shouldered

point typically Upper Solutrean in Iberia (Cortes et al. 1977). Subsequently, Abbé Jean

Roche excavated the Upper Paleolithic deposits at the end of the passage and room 1974

to 1987 (Figure 5.39).

Roche’s excavations revealed 12 stratigraphic levels (Figure 5.41) that he interpreted

in preliminary reports (Roche 1979, 1982). His archaeological interpretation was based

only on the typological and technological analysis of lithic material. Radiocarbon dates

were never obtained despite the recovery of numerous charcoal samples in every

occupation level. Nevertheless, Level 3 was considered Epipaleolithic and Levels 4-9,

Upper Paleolithic. Levels 8 and 9 were classified as Magdalenian based on the presence of

sagaies. Roche (1982) interpreted the association of two human teeth with a cache of pierced

marine shells, lynx canines, red ochre and charcoal as an intentional human burial. He

then interpreted Level 9 as a prepared Magdalenian “floor.” Faunal remains from the

Upper Paleolithic levels included red deer, ibex, horse, wild boar, lynx, small rodents,

small carnivores, birds, and fish. Despite years of excavation and deep soundings, Roche

never reported finding evidence of a Solutrean occupation as suggested by the earlier


Zilhão (1997) re-analyzed the lithic assemblages and concluded that Levels 8 and 9

were Solutrean based on the presence of a few bifacial trimming flakes in the collections

from the 1960s. This was supported by a reinterpretation of the stratigraphy and

consideration of the pointe à cran (Table 5.10).

Suão Caldeirão Physical Archaeology Chronology

Levels  Level A/B/C/Ea loose mixed soils Neolithic &  bp
Bronze Age
Discontinuity Discontinuity Neolithic  bp
intrusions into
Level Eb Loose brown Magdalenian  bp
sandy sediment
with limestone
Discontinuity Discontinuity Perturbation of   bp
and stalagmitic the subadjacent
crust formation levels
Levels  Levels FaFc compact reddish Upper Solutrean   bp
sandy clays with
limestone blocks

Table : Stratigraphic interpretation of Suão and Caldeirão Caves

(After Zilhão 


Figure  : Plan of Lapa do Suão


Figure 5.40: Lapa do Suão

Figure 5.41: Stratigraphic profile of Lapa do Suão (Corte 1, 1974)
(after Roche 1982)





Zilhão further argued that the Solutrean occupation was located mainly near the cave

entrance since the pointe à cran was found there. He concluded that the Solutrean materials

in the back of the cave must have been redeposited since the lower deposits are marked

by a dip. These deposits filled the SW corner of the cave, leveling the area in which the

subsequent Magdalenian occupations took place.

In 2000 and 2001, the faunal assemblages were studied by myself and Maria João

Valente of the Universidade do Algarve. I studied the Magdalenian and Epipaleolithic

levels 3-7 and Valente studied levels 8 and 9 (the supposed Solutrean levels). Preliminary

results were presented in a joint paper (Haws and Valente 2001).

The materials were kept for several years at the Universidade de Porto where Roche

worked. In the late 1980s, the collections were moved to a small regional museum in

Bombarral, a municipality located near the cave. These materials are on loan to the National

Museum of Archaeology in Lisbon where the analysis took place. Unfortunately, no fauna

or charcoal was marked Level 3, the lone Epipaleolithic occupation. Also, many bags

from upper levels lacked source tags, forcing us to leave several aside. As a consequence,

the majority of the material came from Levels 7, 8 & 9.

Charcoal samples were sent to Geochron Laboratories for conventional radiocarbon

dating. This was funded by a Geochron graduate student research award. Although

the dates are not in sequence all are Pleistocene and fall within the Portuguese

Magdalenian between 16,000 and 10,000 years bp (Table 5.11).


Archaeological level Radiocarbon date Lab number

Level    /  GX

Level    /  GX
  /  GX


Level  /  GX

Level    /  GX

Table : Absolute dates for Lapa do Suão
(uncalibrated radiocarbon years bp)

Prior to dating, the charcoal samples were identified by Marjeta Jeraj and myself at

the University of Wisconsin-Madison. The work is still in progress, however the bottom

two levels are complete. Levels 8 and 9 both contain wood charcoal from species comprising

a mixed Atlantic and Mediterranean vegetation cover. These include deciduous oak, pine,

juniper, Rosaceae (wild fruit trees), olive, and other Mediterranean varieties. The Suão

charcoal is similar in composition to the Magdalenian assemblage identified by Figueiral

at Cabeço de Porto Marinho, with the exception of evergreen oak and wild strawberry.

The microfauna from all levels are a typical Mediterranean suite including Microtus

lusitanicus, Arvicola terrestris and Eliomys quercinus. If one accepts the Level 8 date of c.

15,000 bp, then the charcoal and microfaunal data fit the notion that the post-LGM warming

trend was well underway in Portugal prior to the Dryas I.

Faunal analysis for Lapa do Suão

The large mammal remains total 339 specimens of which 83 were identified to species

or genera (Table 5.12). The majority of these are red deer, followed by Iberian lynx (Lynx

pardina) and wild boar. Additional species include fox, wolf and unidentified mustelid

(cf. Meles meles), equid and caprid. As Table 5.12 shows, Levels 4, 5 and 6 contained few

specimens. In contrast, Levels 7, 8 and 9 were much richer in the number and variety of


Level  Level  Level  Level


Red Deer     

Wild Boar  


Table  : Macrofauna NISP in Lapa do Suão

Little can be said of the upper levels due to the scarcity of faunal remains. A burned

and cut-marked Iberian lynx ulna was recovered in Level 6. This might be interpreted as

evidence for the use of lynx for pelts or teeth but it could have been for food. Though lynx

are rare in Portugal today, they were hunted and eaten by rural people until the 1940s

(Valente, pers. com.).

The Level 7 herbivores include both adult and juvenile red deer and wild boar. Only

one burned portion was recovered and four fragments had cut marks, three of them limb

shafts. Carnivores are represented by lynx, wolf, fox and a mustelid. In this level, no cut

marks were found on the lynx.

Levels 8 and 9 contained two adults and one juvenile red deer, a single wild boar,

horse and ibex. As in the previous layer, burned bone is rare. Cut marks were observed

on red deer and wild boar bones, plus a few indeterminate specimens. Most of these are

shallow, oriented diagonal or perpendicular to the longitudinal axis of the bone and likely

from defleshing the carcass of the animal.


Red Deer Level Level Level Level Level



Table  a:Red deer NISP in Lapa do Suão

Table  : Lapa do Suão Rabbit MNE











Some of the wild boar, fox and indeterminate specimens exhibit tooth marks consistent

with carnivore gnawing. In all three levels, some limb shafts show impact fractures and

other breakage indicative of marrow extraction. One red deer phalanx was split

longitudinally as at Picareiro. While carnivore activity may have impacted the assemblage,

density-mediated attrition is unlikely given the near-complete absence of ungulate teeth.

These observations led Haws and Valente (2001) to believe humans deposited the majority

of large herbivore remains, possibly even the lynx.

The NISP for the macrofauna here are so few in number that analyses comparable to

Picareiro Level F are impossible (Table 5.12a). If Magdalenian hunters were bringing

significant numbers of red deer and wild boar into the cave, they only minimally processed

them and transported entire carcasses away. Because Suão is located in a narrow valley

that opens onto a broad coastal plain, residential base camps may have been located close

enough to obviate the need for extended occupations in the cave. This may explain why

so few large animal remains were recovered.


As with Picareiro, rabbit remains numerically dominate the Suão assemblage. Over

5,000 rabbit bones were recovered from Lapa do Suão, representing at least 234 individuals

(Table 5.13). Five hare bones were also identified.

Hockett (1991, 1994, 1995, 1996, 1999) has observed bone damage and skeletal element

patterns typical of raptors, carnivores and humans. A general discussion can be found in

Hockett and Haws (2002). His actualistic studies of raptor nests show distinct, regular

types of damage to rabbit bones by owls and eagles. The eagle owl (Bubo bubo) commonly

nests at the entrances of caves and above rockshelters in Iberia. Hockett (1995) found a

predominance of hindlimbs over forelimbs in owl assemblages. Eagle owl assemblages

in Spain display considerable variability (Sanchis Serra 2000). Some lack forelimbs others

contain more forelimbs than hindlimbs. Cranial elements are more common in some than

others. Lumbar vertebrae and sacra often greatly outnumber cervical and thoracic

vertebrae. The assemblages studied by Sanchis Serra lack abundant foot elements but

that may be due to collector bias. High degrees of breakage are typical of owl assemblages.

Damage typically occurs on the limb epiphyses (especially the greater trochanter of the

femur), vertebrae and innominates (primarily near the acetabulum) (Hockett 1991, 1994,

1995; Sanchis Serra 2000). Spiral fractures are common but bone cylinders are rare. Broken

limbs are generally characterized by epiphyses with attached shaft portions. Beak and

talon punctures occur on 2-3% of bones, usually on one side (Hockett 1991, 1995).

Rabbit bones from carnivore dens and scats typically show signs of etching and

polishing. Large carnivores such as wolves and coyotes completely destroy rabbit bones

during consumption (Schmitt and Juell 1994). Small carnivores often leave bones intact

but with characteristic damage such as tooth punctures on both sides limb bones (Hockett

1989, 1995; Pérez Ripoll 1993; Valente 2000). In Iberia the two main predators of rabbits

are the Iberian lynx and fox. No actualistic studies of rabbit consumption patterns by

Iberian lynx have been undertaken. Sanchis Serra (2000) conducted a study of rabbit

bones from three zones within a single fox den. Hockett (1999) analyzed the bone

assemblage from a carnivore den at the entrance to Picareiro. Punctures occurred on less

than 1% of the total assemblage. These were located on both sides of limb epiphyses and

the innominate. Valente (2000) studied the faunal remains including rabbit from the Early

Upper Paleolithic site, Pego do Diabo. This site contained a carnivore-accumulated

assemblage that was thought originally to have been associated with lithic artifacts (Zilhão


Human-created rabbit bone assemblages are known from definite cultural contexts

within archaeological sites in the Great Basin of the United States, the Spanish

Mediterranean Region and now Portugal (Vila et al. 1985; Drews and Schmitt 1986; Hockett

1989, 1991, 1992, 1995; Schmitt 1990; Rowley-Conwy 1992; Pérez Ripoll 1992, 1993; Hockett

and Bicho 2000; Hockett and Haws 2002). Studies of these sites reveals a great deal of

similarity in skeletal element patterns when humans process and consume rabbit carcasses.

Three sites in the Great Basin all had high representation of mandibles, tibiae, scapulae,

skulls and ribs (Hockett 1994, 1995) (Table 5.14). Vertebrae and sacra were present in very

low frequencies. The long bones, mainly tibiae, were processed for marrow

Element Mineral Hill Cave Hogup Cave NY 




vertebra NISP/MAU   
NISP Total    
MNE Total   
Table : Comparison of rabbit MAU between three Great Basin sites
(After Hockett and Haws  )

extraction by breaking or biting off the epiphyses creating cylinders through which marrow

could be pushed through or sucked out. The tibiae in the rabbit is the largest limb bone

and thus contains the most marrow. This explains why tibia cylinders occur in greater

frequencies in archaeological sites than other limb bones. Ethnographic observations of

Great Basin Shoshone groups by Steward led Hockett (1995) to conclude that the paucity

of vertebrae and sacra at archaeological sites was due to bone meal processing. The

Shoshone regularly pounded and ground axial elements and soft greasy epiphyses on

milling stones.

Tibiae cylinders have also been recovered in large numbers from Upper Paleolithic

sites in Iberia (Vila et al. 1985; Pérez Ripoll 1993; Hockett and Bicho 2000; Hockett and

Haws 2002). At Cingle Vermell in Catalunya, Faro noted an overwhelming majority of

rabbit tibiae were fractured near both epiphyses (Vila et al. 1985). Though Faro does not

suggest marrow removal, the pattern implies the creation of tibia cylinders. Fractures at

the distal ends of the radius, ulna and tibia were interpreted as evidence for the removal

of the feet. Pérez Ripoll (1992, 1993), working independently (and seemingly unaware of

Hockett’s work) came to very similar conclusions as Hockett concerning rabbit butchery

patterns. At the Magdalenian sites, Santa Maira, Tossal de la Roca and Cueva de Nerja, he

attributed the distinct pattern of rabbit long bone cylinders to marrow removal.

Magdalenian and Epipaleolithic hunters created numerous shaft cylinders on the tibia,

femur and much less frequent ones on the humerus (Pérez Ripoll 1993).

Additional taphonomic traces of human agency in rabbit bone assemblages are the

presence of cutmarks and burning. In his analysis of the bones from Santa Maira, a Late

Upper Paleolithic site in southeast Spain, Pérez Ripoll (1993) found up to 60% of the bones

had identifiable cutmarks. There is, however, considerable variability in cutmark

frequencies (Hockett and Haws 2002). Evidence of burning on the feet and distal ends of

lower limbs is also common in archaeological rabbit bone assemblages (Hockett 1991;

Pérez Ripoll 1993). Only 3-6.5% of the rabbit bones from Cingle Vermell were burned.

Though distributed evenly across skeletal parts, many distal tibiae and radii had been

burned. Conversely, at Picareiro, phalanges, metapodials, calcanei and astragali were the

most common element burned indicating whole carcasses were probably roasted with the

feet still attached (Hockett and Bicho 2000).

The majority of rabbit specimens from Lapa do Suão are mandibles, limb bones, in

particular tibiae and femora, and innominates. The abundance of these elements is not

due to differential transport or raptors, which often deposit greater numbers of hind limbs

than fore limbs as shown above. Most of the radii and ulnae are broken in half and distal

humeri, so common in Picareiro, are infrequent. The absence of small elements is probably

due to the use of a wide mesh screen during the excavation as well as curation bias. Table

5.15 shows MNE, MAU and %MAU of rabbit in Picareiro Level F. In Table 5.16 it is clear

that there is a strong bias against small skeletal elements at Suão. This is almost certainly

due to the excavation and screen methods used at each site. At Picareiro, elements as

small as phalanges were piece-plotted. All other small elements were collected in the

fine-mesh screen. Figure 5.42 compares rabbit %MAU in Suão with Picareiro. There are

important differences between the two sites. At Suão, both Level 7 and 8+9 have remarkably

similar body part profiles. The small foot bones aside, Picareiro has better forelimb

representation. Both sites lack axial elements except the pelvis. Hockett and Haws (2002)

compared vertebra representation between accumulators. Human-created assemblages

typically lack vertebrae compared to raptors and carnivores. It would appear that

prehistoric people at both sites ground the vertebrae, sacra and ribs into bone meal as

Hockett (1994, 1995) has noted in the Great Basin and at Picareiro (Hockett and Bicho


Alternatively, density-mediated attrition due to chemical weathering or carnivores

may have played a role in the Suão assemblage formation. Pavao and Stahl (1999) used a

photon absortiometer to measure bone density in several species of leporids. They made

two calculations: one based on Lyman et al.’s (1992) methodology for marmots which

normed volume density using squares or rectangles, and their method of shape-adjusting

using circles, triangles, etc. to reduce air space in the estimation of volume density (Pavao

Table : Rabbit skeletal element representation

at Picareiro Level F
Level F







Suão Level







Table : Skeletal element representation of rabbit at Lapa do Suão

Figure  :Rabbit MAU


Suão  Picareiro F

and Stahl 1999). Though positive, significant correlations were found between their shape-

adjusted VD (VDSA), the scan site density rank shows some striking differences between

the two methods. The results using the Lyman et al. (1992) method match those expected

from other taxa. That is to say, limb epiphyses are generally the weakest structurally and

limb shafts are strongest (Table 5.17 and 5.18). Counterintuitively, however, the VDSA

results show that some of the weakest portions in the traditional method are strongest.

For example, the proximal humerus epiphysis ranks as one of the densest scan sites, higher

than the distal epiphysis which is often one of the strongest limb epiphyses. Both methods

show the proximal tibia is denser than the distal end. The densest portion using the

traditional method is the femoral midshaft, whereas it ranks 10th in the VDSA method.

Figure 5.44 shows there is no clear pattern between VDSA and limb portion representation.

Figure 5.45 shows rabbit limb MNE for Suão Level 7. It is clear that the rabbit limb portions

most under-represented are those traditionally thought of as the weakest structurally.

Therefore, it would appear that something in the shape-adjusted method is badly skewing

the results (Hockett pers. com.). Given the absence of small, usually dense foot bones due

to screen size, full comparisons between skeletal element representation and bone density

using either method cannot be made.

With regard to carnivores, Walter (1984) documented the tremendous destructive

action by dogs of small game bones in hunter-gatherer sites in Australia. However, the

sheer numbers of bones left suggests medium/large carnivores did not impact the rabbit

assemblage. The lack of physical traces such as thinning and polishing, and element

survivorship effectively eliminates carnivores as agents of post-depositional destruction

: Rabbit bone density values
(Adapted from Pavao and Stahl )
VD (LD/BT) rank VD (SA) rank














VD (LD/BT) rank VD (SA) rank






Table  : Top ten rank of rabbit scan sites

Note the femur midshaft rank (FE )


LS Level 7 rabbit

portion count (MNE)







0 .1 .2 .3 .4 .5 .6 .7
Figure 5.44: Lapa do Suão rabbit MNE vs bone density
Density data from Pavao and Stahl (1999)

Figure : Suão

rabbit limb portion MNE


hu ra ul fe ti

pepi pdia dia ddia depi


(Payne and Munson 1984; Schmitt and Juell 1994).

Additional evidence of human consumption is the high degree of fragmentation of

the appendicular rabbit bones. In contrast to raptors and humans, carnivores often deposit

a high number of complete or near complete appendicular elements. At Suão only about

1% of the humeri, femora and tibiae are complete. Both Valente and I observed a number

of proximal and distal long bone fragments but shaft cylinders and fragments (about 60%)

were more common (Figures 5.47 a & b).

These data support idea that the majority of breakage occurs on the meat-bearing

extremities to facilitate the removal of the marrow as observed by Hockett at Picareiro

(Hockett and Bicho 2000) and Pérez Ripoll (1992, 1993) in the Spanish Mediterranean


Valente (n.d.) also studied the bones for cutmarks in order to confirm human agency.

She concluded that about 10% of long bones had cut marks in Levels 7 and 8+9. Most of

these were tibiae, normally on the ventral side of the proximal diaphysis. On these and

other long bones, the marks are normally diagonal to the longitudinal axis, with some

deeper perpendicular marks. Cutmarks on the mandible are diagonal to the anterior/

posterior axis. She interpreted these data as evidence for skin removal rather than

disarticulation or fracturation, which may have produced deeper marks.

Based on work by Hockett (1991, 1994, 1995, 1996, 1999), humans likely deposited

most of the rabbits. Only a few modifications by raptors and small carnivores were

observed. These include a few bones with multiple and single punctures. How were

rabbits procured? There is no evidence for nets or cordage from any Upper Paleolithic

Figure 5.47a: rabbit tibiae cylinders from Lapa do Suão Level 7

Figure 5.47b: rabbit femur cylinders from Lapa do Suão Level 7

site in Portugal. However, this technology is known archaeologically from at least the

Gravettian in central Europe if not much earlier in other parts of the Old World (Adovasio

et al. 1996). There is little doubt that all Upper Paleolithic peoples utilized fibrous material

to makes nets and cordage useful in traps and snares. Rabbit drives are known

ethnographically from the Southwest and Great Basin of the United States (Shaffer and

Gardner 1995). Schmidt (1999) used element representation in rabbit assemblages from

Arizona to determine whether or not drives were used prehistorically. One site, the Five

Feature site, contained high NISP and overall percentage of rabbit bones in the faunal

assemblage. Interestingly, the elements represented are distal tibiae, calcanei, astragali,

tarsals, metapodials and phalanges. Only two bones out of 802 came from another skeletal

element. This patterning led Schmidt (1999) to conclude that the Five Feature site was the

processing location for rabbits collected in a prehistoric communal drive. If this element

representation pattern can be seen as diagnostic, then neither Picareiro nor Suão would fit

into this category. Further evidence suggestive of rabbit drives in prehistory is provided

by Hudson (1994). At one site in California (KER-526) she noted that cranial elements far

outnumbered limbs and other parts. Hudson (1994) concluded that large numbers rabbits

had been taken through drives with the heads subsequently removed prior to transport.

A few were probably consumed onsite accounting for the additional elements. This pattern,

if considered diagnostic, is also not apparent at Picareiro or Suão.

Considering rabbit ecology, there are behavioral differences between the European

rabbit and hares that probably make drives unlikely for the former and more likely for the

latter (Hockett 1992; Hockett and Haws 2002). The European rabbit is a territorial animal

that forms harems and lives in burrows (Garson 1981; Soriguer and Rogers 1981). They

are best hunted by stalking individuals, trapping and snaring, or digging them out of

their holes. Because rabbit warrens are easily recognizable to the trained eye, prehistoric

hunters would have known their locations and regularly exploited them. Hares, on the

other hand, are solitary and live in nests on the ground surface. Hare hunting would

probably have been less profitable than rabbit because they would be too infrequent to

hunt individually. They do, however, congregate and migrate in large numbers during

stressful times (Angerman 1981). Hunters could much more easily drive large numbers

of hares into nets at various times of the year (Hockett and Haws 2002).

Rabbits at Picareiro and Suão were more than likely taken by traps , snares and possibly

with bow and arrow (Hockett and Bicho 2000). Net hunting was unlikely due rabbit

behavior (Lupo and Schmitt 2002). In all likelihood, rabbits were hunted by individuals

or small groups who set out traps or snares while engaged in other activities. As with

Picareiro, numerous rabbits were stripped of meat and probably consumed onsite. The

fact that marrow was removed from the humeri, femora and tibiae provides further

evidence of immediate consumption. It is very likely that additional carcasses, possibly

smoked rabbit meat and axial parts of the skeleton were transported to other locations



Several species of birds were present in the Upper Paleolithic levels. A total of 118

bones were recovered in Level 7 (Table 5.19). The most representative species are Anas

platyrynchos (mallard) and Alectoris rufa (red-legged partridge). No remains of raptors


were found. Some of the duck bones have cut-marks, mostly on the humerus, coracoid

and ulna (Figure 5.48). According to Bochenski et al. (1999) golden eagles and other raptors

that prey on other birds, leave higher percentages of sterna,

Table : Bird NISP at Lapa do Suão

Level  Level
Alectoris rufa
Anas platyrhynchos
Corvus monedula
Garrulus glandarius
Pyrrhocorax pyrrhocorax
Tordus sp

coracoids, scapulae and humeri in the unswallowed portion than other elements. Owls

typically leave higher proportions of wing elements. At Suão, the most frequent bird

bones are humeri, radii, coracoids, carpo-metacarpi and ulnae. The latter are usually

missing both epiphyses forming bone cylinders though not for the same reason as those

from rabbits. Bird bones do not contain marrow. Instead they are hollow to aid in lift and

flight. In this case, the missing epiphyses are probably due to eating habits whereby the

soft epiphyses are chewed off (cf. Steadman et al. 2002). Still, the possibility for bird bone

stock is open for suggestion. Garcia Petit (1995) has argued for such a practice during the

Magdalenian at Bora Gran in Catalunya. Several limb elements from geese and bustard

were recovered as bone cylinders. A similar pattern is documented at the Magdalenian

site Grotte de les Églises in southern France (Laroulandie 1998). Lefèvre (1992) observed

this phenomenon in prehistoric and historic sites in Patagonia. She attributed the lack of

epiphyses to human chewing behavior eased by incomplete fusion. Raptors or carnivores


Figure 5.48: cutmarked duck coracoid

cannot be ruled out as agents in the deposition of bird bones at Suão but the element

representation, presence of cutmarks and breakage patterns strongly suggests

predominately human agency.

Aquatic fauna

As for the marine fauna, Roche (1982) reported a variety of molluscan species, clams,

mussels, limpets and gastropods and a single fish, gilthead (Table 5.20). The marine shells

were almost certainly all ornamental as there was no shellmidden reported and the number

of shells is low. Many are perforated, especially the Littorina and Nassarius. Nevertheless,

the collection of these species, mostly from littoral environments, a fairly large exploitation

territory for Suão. Given the screen size used, the fish remains may be greatly under-


Table  : Marine and freshwater shellfish at Lapa do Suão

Level  Level
Cerastoderma edule
Mytilus edulis
Scrobicularia plana
Solen marginatus
Tapes decussata
Patella sp
Cerithium vugatum
Littorina obtusata
Nassarius reticulata
Trivia monacha
Turritela sp
Semicassis undulata
Theodoxus fluviatilis


To summarize, the Suão fauna is numerically dominated by rabbit. Red deer, wild

boar, equids, caprids and carnivores are also present. Additional small animals include

birds, molluscs, fish, reptiles, bats, voles and dormice. Given that most of the cave

sediments were excavated it is likely that the existing sample of Magdalenian artifacts

and food refuse represents the nature of site function. However, the possibility that many

of the animal bones were discarded outside the entrance of the cave could mean that the

sample from the interior is strongly biased in favor of smaller, more fragmented remains.

The preponderance of rabbit fits the general pattern in Upper Paleolithic caves and

rockshelters. For ungulates, red deer is most abundant, as it is in the Magdalenian levels

at Caldeirão and Picareiro. The absence of auroch is perhaps strange since Suão is located

much closer to prime auroch habitat than Picareiro. The fauna, wood charcoal and

radiocarbon dates conform to Roche’s conclusion that the site is Magdalenian and the

Solutrean artifacts derive from an occupation that eroded away. That Roche ignored the

numerous rabbit bones in his publication may be due more to the fact that most of these

were found in later, unpublished excavations than a dismissal that humans hunted rabbits.

Birds are relatively rare in Magdalenian sites in central Portugal. Suão and Caldeirão

are the only sites where waterfowl and partridges, common prey types in Spanish

Magdalenian sites, have been found. Picareiro has a few bird bones but these are of

songbird size (Bicho et al. 2000). Suão is located upslope from a small stream winding

through a narrow valley. Within a kilometer or two, this valley opens to a low coastal

plain. This is a fairly unique location for a fauna-bearing Late Upper Paleolithic site in

Portugal and perhaps explains the presence of several avian prey types. However, it may

also relate to differences in the season of occupation of Suão compared to other sites further


Seasonality determination at Suão is made difficult because of the absence of teeth

for sectioning or crown height measurements. The primary evidence lies in the rabbit

mortality profile. The rabbit assemblage is dominated by adults but does contain a few

juveniles, unlike Picareiro which is almost entirely comprised of adults. Of a total of 2,444

rabbit elements in Suão Level 8+9, Valente (n.d.) identified 94 juvenile rabbit elements

from approximately 13 individuals, about 9% of the total MAU. At Picareiro, Hockett has

reported that 99.4% of the rabbit limb epiphyses were fused (Hockett and Bicho 2000).

Rabbit seasonality at Picareiro was determined by comparison of the mortality pattern

and modern rabbit ecology. Rabbits in Spain and Portugal have two birthing peaks

coinciding with fall and spring rainfall (Soriguer and Rogers 1981). There is a high mortality

rate for juvenile rabbits so that within a few months the juveniles have either matured or

been killed by predators. Therefore, archaeological assemblages dominated by adult rabbits

may have been formed during winter and/or summer (Hockett and Bicho 2002). Unless,

of course, hunters were selectively targeting adults. Lupo and Schmitt (2002) have argued

that traps and snares tend to catch adults more often than not because juveniles either do

not follow adults into traps or are too small to trip snares. If however, one accepts the use

of mortality patterns to determine season of capture, then Suão may have been occupied

slightly earlier or later during the year than Picareiro when subadult rabbits were still

present, perhaps the early Fall or late Spring.

5.5 Additional fauna-bearing Final Upper Paleolithic and Epipaleolithic sites in

Portuguese Estremadura

Gruta do Caldeirão

Much of the data used to reconstruct Late Pleistocene environment and Upper

Paleolithic subsistence in central Portugal comes from Caldeirão Cave. As mentioned

above, Caldeirão (‘cauldron’ in Portuguese) was excavated in the 1980s by Zilhão. The

Middle and Upper Paleolithic macrofaunal assemblages from Caldeirão were recently

studied and published by Davis (2002). This was the first systematic study of the Paleolitic

material and his results form the basis for discussion here.

As with Picareiro and Suão rabbit is the most abundant animal represented. The

rabbit assemblages have been analyzed by Newton (n.d.) but only NISP is available as the

full results are unpublished. The ungulates in the Magdalenian level are best represented

by red deer, followed by wild boar, horse, aurochs, chamois, ibex and roe deer. Carnivores

include lynx, leopard (Panthera pardus), wildcat, fox and badger (Meles meles). Other small

animals include hare, beaver (Castor fiber), birds and fish. The birds are represented by

chough (Pyrrhocorax pyhhrocorax), magpie (Pica pica), pigeon (Columba palumbus), partridge

(Alectoris rufa) and little owl (Athene noctua) (Davis 2002).

The methodology Davis used differs from Americanist zooarchaeological tradition.

He followed the “diagnostic zones” method outlined by Watson (1979) in his study of

Khirokitia. Comparisons between Picareiro and Caldeirão are therefore difficult because

of differences in analytical methods. The diagnostic zones method was designed to avoid

problems associated with counts based on NISP and MNI. Watson (1979) argued that

NISP counting would artificially inflate the skeletal element frequency because the same

bone could be counted more than once if fragmentation was high. Potentially, each

specimen could be assumed to come from a different animal (Grayson 1984). In addition,

MNI estimates would also be inflated if the whole site was not excavated, which is almost

always the case. As Grayson (1984) has also pointed out, MNI counts exaggerate the

relative importance of rare species. To solve these problems, Watson devised a list of

diagnostic zones on each element that would serve as criteria for counting a specimen or

not. The method results in a count that appears similar to landmark MNE.

Davis (2002) used a very restricted diagnostic zone approach. For instance, the tibia

count was based solely on the medial half of the distal epiphysis. So, the lateral half of the

distal end, the entire shaft and the proximal end of the tibia would not be counted. All

specimen counts were based on mandibular teeth and limb epiphyses. As Davis states,

this was done ”so that data from Caldeirão can be easily interpreted and used by other

zoo-archaeologists working on the same material…” (Davis 2002: 33). Furthermore, he

reasons that the selected zones provide sufficient data on body part representation to be

used in determining butchery and transport patterns. From a zoological point of view,

this may be useful in identifying species. However, from an archaeological standpoint,

this method leads to a serious bias in the data. Clearly, this method precludes direct

comparison with Picareiro, especially with regard to limb elements. As shown above,

limbs were divided by portion and the MNE was calculated for each portion. Though

bones with more than one portion are counted in each category they are not “counted

twice.” Each bone specimen is counted and the recorded portions are then tallied to arrive

at MNE estimation. In Davis’ method, a specimen is only counted if a certain diagnostic


part is present. A tibia represented by the proximal end would end up in the “unidentified”

and “non-countable” pile. If every limb specimen was a complete element this method

would be satisfactory. However, this is almost never the case in archaeofaunal assemblages.

Element fragmentation and density-mediated attrition of soft epiphyses due to natural

weathering, carnivore gnawing or human grease processing will result in under-

representation of limb elements (Bunn 1986; Bunn and Kroll 1986; Marean and Spencer

1991; Marean 1995; Marean and Frey 1997; Marean and Kim 1998). The comparison between

Figures 5.31 and 5.31a showed the differences in limb element representation at Picareiro

when limb shafts are counted or not. Counts based solely on one end of a limb bone also

precludes any comparison between element survivorship and bone density. Therefore,

density-mediated attrition, carnivore activity and human butchery and transport of

ungulates at Caldeirão cannot be evaluated. Only 93 specimens or 4% of the Magdalenian

assemblage (N=2164) were identified to element or species using the diagnostic zones


The Caldeirão sample does provide some useful biometric data, which when combined

with similar data from Picareiro, can be compared against the larger data set on Pleistocene

large mammals in Portugal published by Cardoso (1995) (Table 5.21). Further comparison

between Cantabria (Mariezkurrena and Altuna 1983; Klein and Cruz-Uribe 1994) and

Mediterranean Spain (Davidson 1989; Morales 1995) can provide a context for

reconstructing ungulate biomass in central Portugal during the Late Pleistocene. This is

useful to understanding prey selection, resource quality and availability for prehistoric

hunter-gatherers. Davis’ (2002) measurements on Caldeirão red deer shows no size


Table 5.21: Red deer Metatarsal Bd

Mean Max. Min. N
Fontainhas (P) 38.8 43.0 31.5 3
Caldeirão 38.1 38.8 36.4 2
Casais Robustos (P) 42.0 1
Lapa da Rainha (P) 40.0 41.0 39.0 2

Caldeirão Fb 39.2 1
Fa 45.2 48.0 42.3 2
La Riera 44.0 49.5 41.5 4
Altamira 45.4 47.2 44.1 6

Caldeirão Eb 37.5 1
Picareiro F 39.9 44.0 37.4 3
João Ramos (P) 36.2 38.5 34.0 5
La Riera 44.6 47.0 41.0 4
Urtiaga 46.4 52.5 42.7 15
Tito Bustillo 44.7 49.5 42.0 17
La Paloma 43.6 48.5 40.5 25
Altamira 45.1 49.2 42.5 14
El Castillo 44.4 53.0 39.6 61
El Juyo 44.7 48.9 41.4 25

Postglacial 41.7 50.5 35.4 23

Modern 36.1 41.0 33.0 29
Data from Cardoso (1995), Mariezkurrena & Altuna (1983)
Klein & Cruz-Uribe (1994), and Davis (2002).. P= paleontological

Table 5.22: Red deer astragalus GLl

Mean Max. Min. N
Middle Paleolithic
Furninha (P) 57.0 63.0 51.0 2
Figueira Brava 58.3 1
Lorga de Dine (P) 56.6 1
Caldeirão 54.7 1

Fontainhas (P) 53.0 56.0 50.0 15
Caldeirão 55.3 1
Pego do Diabo (P) 54.0 54.3 53.8 2
Gruta de Salemas 51.7 52.0 51.7 2
Pedreira de Salemas 53.5 58.7 48.0 9
Lapa da Rainha (P) 54.0 1

Parpalló 6 50.8 54.7 44.6 33
7 49.6 52.7 46.8 6
9+10 50.6 54.1 45.9 24
Algar de Cascais 58.0 1
Escoural 50.4 54.5 46.0 8
Caldeirão 54.4 57.7 52.7 4

Parpalló 1 51.8 57.2 46.2 47
2 51.8 55.6 47.9 36
3 49.9 55.2 45.7 30
Bora Gran 57.1 12
Urtiaga 57.2 67.1 50.7 22
Nerja 53 1
Caldeirão 52.8 54.3 50.7 4
Picareiro F 53.1 54.9 50.7 5
João Ramos (P) 54.2 55.0 53.5 2

Data from Davidson (1989), Morales (1995) and Davis (2002)..

P= Paleontological

differences between the Middle Paleolithic and the end of the Pleistocene. Size diminution

in red deer appears to have occurred sometime during the Holocene when the data are

compared to modern red deer from Navarra (Mariezkurrena and Altuna 1983; Davis 2002).

The red deer from sites in Portugal are smaller on average than those from Cantabrian

Spain at least during the Early Upper Paleolithic and Magdalenian. The only EUP sample

from Spain is the El Castillo sample (not shown) that probably dates about 15 ky earlier

than the OIS 2 Portugal sample. This group was much larger (X=47.7: Klein and Cruz-

Uribe 1994: Fig.5) than the specimens from Portugal. During the Solutrean the Caldeirão

specimens are within the range of the Cantabrian ones except the sample from Fa. Level

Fa at Caldeirão dates after the Last Glacial Maximum and may be related to the so-called

“Solutreo-Gravettian” from Mediterranean Spain (Zilhão 1997). Does this mean that LGM

forage quality was similar between Cantabria and Portuguese Estremadura? Zilhão (1990,

1997) has argued on a number of occasions that Estremadura had cool dry Artemisia-

steppe conditions during the LGM. Comparing the Magdalenian samples it would appear

that Cantabria may have had better red deer forage than Estremadura after 15,000 bp.

This corresponds to a return of warm, humid mixed Mediterranean evergreen and

deciduous forests in Portugal which may have lowered the potential for red deer body


No red deer metatarsal measurements are available from Mediterranean Spain but

Davidson (1989) and Morales (1995) report astragali dimensions from Parpalló, Urtiaga,

Bora Gran and Nerja. These are shown with measurements from Picareiro, Caldeirão

(Davis 2002) and other Upper Pleistocene sites in Portugal in Table 5.22. Once again the

Cantabrian samples are much larger. Five astragali from Picareiro fall easily in the range

(on the larger end) of those from Parpalló. They also fall in the same range as the Caldeirão

sample. Davis (2002) argued that there appeared to be a decrease in astragali size between

the Late Pleistocene and Early Holocene. The Holocene samples do appear to be smaller,

however those samples are still in the range for Parpalló. It could be that the Parpalló

sample has a larger number of females and is thus skewing the range. Mariezkurrena and

Altuna (1983) documented sexual dimorphism in red deer from Navarra though there

was overlap. The largest of the modern Navarra sample is smaller than the smallest one

from Caldeirão. It could be that males were represented by the astragali from Picareiro

and Caldeirão.

Bocas, Casal Papagaio, Pena da Mira

Bocas rockshelter was excavated in the 1930s by Manuel Heleno, an historian with

the National Museum. The site had several components including Upper Paleolithic and

Epipaleolithic but they were never reported. Heleno did not keep field notes and the only

clue to the site comes from his drawing of the stratigraphic profile. Bicho (1995-1997)

recently analyzed the lithic material and obtained radiocarbon dates on bone and shell for

the lower levels. The Final Magdalenian and Epipaleolithic levels date 10,110-9,900 bp.

The small faunal assemblage includes auroch, horse, red deer, wild boar, ibex and chamois

and is being studied by Valente. In addition to the terrestrial fauna, a small shellmidden

was excavated though the details are too sketchy to know the exact nature of the deposit.

Gruta da Casal Papagaio is a cave located approximately 10km north from Picareiro

near Fátima. This site was excavated in the 1980s but only a preliminary report has ever

been published (Arnaud and Bento 1988). Arnaud and Bento (1988) report two radiocarbon

dates of 9,710bp and 9,650 bp on marine shells. The site contained the remains of two

humans. Numerous shells, some perforated, including cockles, clams, mussels and even

crab were excavated. Terrestrial fauna included red deer (antler), caprids, rabbits, rodents

and fox. Though charcoal was recovered, no hearths or other features were found. No

lithics or ceramics were encountered either. The cave lies 35-40 km inland from the Atlantic

Ocean. During the period of use, the sea was about where it is today, possibly slightly

further away.

Pena de Mira is a large rockshelter located on the south face of a scarp along the

Polje of Minde (Figures 5.49, 5.50, 5.51, 5.52, 5.53). This large sinking basin lies only 2 km

west of Picareiro. Most of the intact sediment was removed during the construction of a

large cistern over a natural rainy season spring. A small section remains and was sampled

by STEA for the Carta Arqueologica de Parque Natural das Serras d’Aire e Candeeiros in

the late 1980s. A radiocarbon date of 8,500 was obtained on charcoal (Zilhão and Araújo

1991). The site contains bone fragments and marine shells but they are encased in a breccia

of limestone éboulis.

Figure 5.49: Pena de Mira rockshelter


Figure 5.50: The Polje of Minde during winter

Figure 5.51: same view in summer

Figure 5.52: The Polje of Minda with the Serra d’Aire in the background
(Picareiro is just left of center along the top of the mountain, Pena de Mira
isalong the left margin of the water left of center)
Figure 5.53: Pena de Mira in summer

Lapa dos Coelhos

Several Paleolithic sites are know along the limestone scarp known as the Arrife d’Aire.

This lies against the southeast face of the Serra d’Aire on the opposite side from Picareiro.

Where the source of the Almonda River flows out of the limestone, there are a series of

caves dated to the Lower, Middle and Upper Paleolithic. One of these, Lapa dos Coelhos,

has thick Magdalenian deposits that have recently been excavated by Francisco Almeida.

The faunal assemblage has not been fully analyzed and published so only preliminary

indications can be discussed. As implied by the name, this site was full of rabbit bones.

Approximately 1,000 rabbit bones had been identified after the 2000 field season (Hockett

and Haws 2002). No details on the assemblage composition have been published.

Buraca Grande & Buraca Escura

Two Upper Paleolithic sites, Buraca Grande (Big Hole) and Buraca Escura (Dark Hole),

are located along a tributary of the Mondego river in the limestone uplands of the northern

part of Estremadura. Whether or not this are was part of the same exploitation territory as

the people who used the caves 50 km to the south is a matter of debate. These caves were

recently excavated and only preliminary reports have been published (Aubry et al. 1997;

Aubry et al. 2001). Buraca Grande has occupations attributed to the Gravettian, Proto-

Solutrean, Solutrean followed by a possible erosional event. Hints of a Magdalenian

occupation were suggested by an accelerator date of 13,050 +/- 100 bp on a decorated

bone (Aubry et al. 1997). Of interest is the microlithic level attributed to the early Mesolithic

with three dates between 8,680 +/- 40 and 7,580 +/- 30 bp. Aubry et al. (1997) report the

faunal assemblage is dominated by lagomorphs with some evidence of burning and

anthropic fracturing. In addition, red deer and wild boar were identified. The bones have

been analyzed by Jean-Philip Brugal but nothing has been published to date. Marine and

estuarine shells were also found but no indication of a midden was reported. Buraca

Grande lies about 40 km inland from the Atlantic, roughly the same distance as the shell-

bearing sites mentioned above. Although the species are of ornamental type, their presence

suggests some movement between the coast and interior.

Buraca Escura is located across the valley from Buraca Grande. This cave has Middle

and Early Upper Paleolithic deposits but is noteworthy because of the near absence of

lagomorphs. Wild boar is also absent and red deer is less abundant than ibex, horse and

aurochs Aubry et al. (2001). As with later cave use in Portugal, large game counts are still

quite low. The charcoal assemblage is dominated by pine (Pinus sylvestris), boxwood

(Buxus sempervirens) and Leguminosae pointing to either a cooler climate or reflecting

available vegetation on the north face of the Valle do Poio Novo. The absence of rabbit

and wild boar is probably due to the unfavorable habitat near the cave during its occupation

since these animals are known from Gravettian contexts to the south at Anecrial (Hockett

and Haws 2002).

5.6 Discussion

Comparisons between the archaeofaunas from the caves and rockshelters in Portugal

permit a general understanding of the human use of these locations during the Late Upper

Paleolithic and Epipaleolithic. A number of questions arise including (1) what types of

sites do these caves and rockshelters represent?; (2) Are they seasonal?; (3) Do they

represent the overall subsistence economy of Late Upper Paleolithic people in Portugal?;

(4) Is there evidence for resource depression and a subsequent widening of diet breadth?;

(5) How does the Late Upper Paleolithic and Epipaleolithic of central Portugal compare to

Cantabria and Mediterranean Spain.

There appear to be some important similarities between Picareiro, Suão and possibly

Caldeirão with regard to the processing of rabbit. Whole carcasses were roasted and

consumed. Marrow was systematically and regularly removed from the limbs. The

macrofaunal assemblages differ quite strikingly at first glance. Almost nothing can be

said of Suão regarding ungulate carcass use. Elements of red deer and wild boar were

deposited in higher frequency than equids and caprids but the sample is extremely small.

Both Picareiro and Caldeirão have macrofaunal assemblages dominated by red deer and

wild boar during the Magdalenian. On the other hand, Suão and Caldeirão contain several

carnivore species whereas Picareiro has almost none. Because of the different methods

used, the only comparisons that can be made with Caldeirão are based on raw specimen

counts, taxa present and biometrics.

Archaeologists working in the Great Basin have devised a relative abundance index ,

also called the Artiodactyl Index (AI), to measure the proportion of large to small animals

in a given site (Ugan and Bright 2001). This index is calculated by dividing the number of

large animal specimens by the sum of large and small animal specimens. Using the

commonly held assumption that large game outranks small game, Ugan and Bright (2001)

used the AI as a measure of foraging efficiency. Stiner et al. (2000) also used this index to

discern trends in the percentage of small game in Middle and Upper Paleolithic sites in

the Mediterranean. Davis (2002) used a rabbit:ungulate ratio to show an increase in rabbits

from the Mousterian to the Magdalenian at Caldeirão.

The AI measures between the Picareiro, Suão and Caldeirão are given in Table 5.23.

At first glance, it appears that the ungulate proportion at Caldeirão decreases through

time from the Mousterian to the Magdalenian suggesting a decline in foraging efficiency.

The AI values for Picareiro show almost the opposite trend for the Magdalenian sequence.

Table  : Abundance Indices for sites in Portugal

Ungulate NISP Rabbit NISP Abundance

Index (AI)
Picareiro E upper Magdalenian   
E lower Magdalenian

F Magdalenian   
G Magdalenian 
Caldeirão Magdalenian    


Picareiro has a higher number of identified large mammal specimens so the index is higher,

though only slightly. Suão is similar to Picareiro.

Davis (2002) noted an increase in the number of unidentified specimens to identified

ones between the Mousterian and Magdalenian. This increase corresponds to the increase

in the rabbit:ungulate ratio. Thus, the apparent decline in foraging efficiency seen by a

reduction in the AI index through time is an illusion. The AI is simply tracking an increase

in the degree of ungulate bone fragmentation through time. Higher ungulate bone

fragmentation results in fewer identifiable pieces. On the other hand, rabbit bones are

rarely fragmented beyond recognition during human butchery practices so their degree

of fragmentation should remain constant through time. Dry bone breakage, trampling

and marrow cracking can have a major affect on ungulate bone assemblages. Changes in

exposure, site occupation time and time spent processing carcasses can result in different

degrees of fragmentation through time. Measures based on numbers of identified

specimens are therefore unreliable in calculating abundance.

Table 5.24 shows the total number of ungulate and rabbit bone specimens for Picareiro,

Suão and Caldeirão.

Table  : Comparison of macrofaunal and rabbit NISP from Magdalenian

sites in central Portugal
Picareiro Suão Caldeirão
indeterminate ungulate  

Ungulate NISP   
Rabbit NISP 

different counting method used

Total ungulate NISP:MNE ratios can be calculated to measure the degree of bone

fragmentation at Picareiro and Suão. However, estimating bone fragmentation using

NISP:MNE for Caldeirão is impossible because bones were not counted in the same manner.

The number of identifiable specimens is unknown and therefore no reliable MNE estimation

can be made.

total ungulate NISP:MNE

Picareiro F 
Suão (

The difference between these two sites and Caldeirão in Table 5.24 is readily apparent.

Caldeirão has more large animal specimens and roughly equal or slightly fewer rabbits in

the Magdalenian levels. Does this mean that there were greater numbers of ungulates

brought to Caldeirão? Assuming a similar degree of fragmentation it would appear that

Caldeirão actually contained a greater percentage of ungulates than Picareiro. Given the

high NISP:MNE ratio at Suão and Picareiro it is unlikely that the Magdalenian assemblage

from Caldeirão could be significantly more fragmented.

Is the higher percentage of ungulates in the Magdalenian at Caldeirão due to significant

differences in site function? Caldeirão is located further inland than Picareiro but at much

lower elevation and closer to the preferred habitat of many game animals. Therefore it

should be expected to have more taxa and more animals represented. Each site appears to

have been utilized as a temporary carcass processing site. Large game were intensively

processed for meat and marrow and much of it seems to have been consumed onsite at

Picareiro and Caldeirão. At Suão the ungulates were mostly transported away. On the

other hand, all three sites were rabbit carcass processing stations. Large numbers of rabbits

were butched and consumed. There is evidence that many more were transported away

from Picareiro and Suão. The rabbits from Caldeirão have not yet been published so no

definitive conclusions can be made.

5.7 The broader regional context

Comparing these sites with the other poorly known caves and rockshelters in

Estremadura a number of observations can be made. First, it would appear that caves and

rockshelters, while providing most if not all of the direct evidence for Late Upper Paleolithic

subsistence, do not contain the same quantities of large animal bones as they do in other

regions of Europe, especially Cantabrian Spain and Southwest France. This does not mean

that environments were poorer. Large gregarious herds of reindeer, bison and horses

simply did not exist south of the Pyrenees. The animals hunted south of the Ebro River

had different biogeographic and ecological characteristics. Large animals were rare and

medium ungulates were solitary or found in small groups. In contrast, small game was

much more abundant, probably due to the milder climatic conditions of near-coastal Iberia.

Coastal and plant resources were probably more plentiful as well. The opportunity for

high numbers of large mammal bones to accumulate from processing of mass-kills would

not have existed. Instead most of the locations were used for intensive rabbit carcass

processing with occupations lasting long enough to result in the deposition of small but

highly fragmented large mammal assemblages. These sites do not seem to be long-term


Recent syntheses of the archaeological record from the Mediterranean regions of

Spain by Aura et al. (1998) and Villaverde et al. (1998) suggest a similar human land-use

and subsistence pattern to the one in Portugal. As noted in previous chapters, both regions

share a similar climate and environment. Both regions are characterized by intensive

rabbit exploitation during the Upper Paleolithic. Cultural connections between the two

can be seen from at least the Solutrean through formal similarities in projectile points. In

Portugal, Parpalló points were found at Caldeirão, Salemas and Passal (Zilhão 1997). The

‘Solutreo-Gravettian’ of Mediterranean Spain may exist in Estremadura as well (Zilhão


Aura Tortosa and Pérez-Ripoll (1995) show that rabbit often comprises over 90% of

faunal assemblages in Mediterranean Spain during the Upper Magdalenian (MSM-

Magdalenian Superior Mediterráneo 13,000-11,000 bp). Table 5.25 shows rabbit NISP is

23,850 from sites in the region while larger fauna (primarily red deer and ibex) number

approximately 10,000. Most of the rabbit comes from Cova Matutano dated approximately

between 13,960-11,410 bp and Cova de les Cendres levels IX-X dated 12-16,000 bp (Olària

1999; Villaverde et al. 1999). Over half of the Matutano sample comes from Matutano I

(the upper levels) dated 11,410-12,500 bp. Olària reports large hearths in this level which

resemble those from Picareiro F and G. The limb fragmentation is described, “the tibia is

broken in the first moment by the distal shaft, removing the feet whole and without

disarticulating them. In the second phase, this bone is distarticulated from the femur and

broken by the part with the most thin walls, removing the proximal epiphysis” (Estévez

in Olària et al. 1985: 87). This suggests the systematic creation of rabbit bone cylinders to

access the marrow as documented in the Portuguese sites. Large game bones are also

highest in this level and the percent identified is only slightly higher than Picareiro

suggesting a high degree of fragmentation here as well. In Matutano IV, dated 13,370-

13,960 bp, another spike in rabbit occurs where smaller, “pseudocircular” hearths were

found (Olària 1999: 423, Table 2). This situation is similar to Level J in Picareiro where a

small oval hearth yielded a large concentration of rabbit bones. This level is undated and

its interpretation often changes as a result (Hockett and Haws 2002; Bicho et al. 2002; Bicho

pers. com.). Overall at Matutano, the majority of red deer skeletal parts represented are

phalanges, carpals and isolated teeth. Limbs are greatly underrepresented and it is likely

Table 5.25: Rabbit vs. large game NISP in Mediterranean Spain

Magdaleniense Superior Rabbit NISP Large game NISP
Mediterraneo (13-11k bp)
Cova de les Cendres 5139 1166
Parpalló 55 3007
Tossal de la Roca 1561 364
Nerja 1936 2613
Cova Matutano 13738 944
Cova dels Blaus 739 78
Chaves 682 1751
Total 23850 9923

Epipaleolitico Microlaminar
Mediterraneo (11-9k bp)
Cingle Vermell 3814 153
Santa Maira 1186 737
Picamoixons 346 21
Cova dels Blaus 1601 268
Nerja 2425 1025
Les Malladetes 28 66
Tossal de la Roca 1462 545
Total 10862 2815

Epipaleolitico Geometrico
Mediterraneo (9-7.5k bp)
Tossal de la Roca 111 630
Santa Maira 35 67
Cueva de la Cocina 93 661
Cova Fosca 1066 157
Total 1305 1515

Total 36017 14253

Data from Vila et al. (1985), Aura & Pérez Ripoll (1995),
Villaverde & Martinez Valle (1995), Olaria (1999), Aura et al (2002),
Castaños (1993), Davidson (1989), Verges Bosch (1996)

that the unidentifiable remains are mostly limb shafts. The published drawings of fracture

patterns show few epiphyses with attached shafts and only a few shafts identified to

element. In Matutano as well, phalanges were split open to extract marrow. Estévez

shows limb representation rises dramatically when the shaft fragments of the unidentifiable

portion are considered (Estévez in Olària et al. 1985: 91).

Additional sites in the SMR exhibiting a similar usage include Tossal de la Roca

and Cova de les Cendres. Both sites are numerically dominated by rabbit with moderate

large game NISP. In their analyses, Aura and Pérez Ripoll (1995a, 1995b) show a high

degree of ungulate limb bone fragmentation at Tossal de la Roca. Limb elements are

mainly represented by shaft fragments. In Nerja and Santa Maira, ibex limb epiphyses

are nearly absent and the overwhelmingly majority of specimens are midshafts. At Tossal

de la Roca, midshafts comprise about 70% of the ibex limb portions in the Magdalenian

levels in the interior of the shelter. Shafts decrease significantly in the Epipaleolithic levels

of the exterior excavation where distal fragments approach 50% of the element portion.

Levels IIa, I and Sup. from the exterior of the shelter generally do not have a high degree

of fragmentation. The authors attribute this difference to physical factors affecting the

bones and a smaller excavated area. Nevertheless, there may be significant differences

between the two periods. Since limb elements have different bone densities the variability

may be due to differences in the representation of certain elements. For instance, in the

grouped distribution graphs, the Magdalenian levels have a predominance of similar

hindlimb portions, which appears to be femur and tibia shafts (Aura and Pérez Ripoll

1995: Figure 5). In the exterior area, the dominate portion appears to be the distal tibia. It

could be argued that the distal tibia should be present equally in the interior relative to

midshafts. Unfortunately, the percentages of each limb portion are based on NISP instead

of MNE. A higher degree of limb fragmentation in the interior would create greater

numbers of midshaft fragments thus lowering the percentage of distal ends. If MNE was

considered then those effects would disappear and the true pattern would emerge.

Nevertheless, they attribute the absence of limb epiphyses to grease processing, not to

carnivore chewing or density-mediated attrition due to in situ weathering.

The practice of using NISP in skeletal element analyses is the norm in Spanish

zooarchaeology and it makes comparison difficult. As Straus (1987) notes for faunal reports

from the Cantabrian region,

None of the lists separate long bones among proximal, mesial, or distal
elements or give any fragmentation information. It is not reasonable to
attempt to extrapolate minimum numbers of individuals per anatomical
element, given the absence of this information. For these reasons, full inter-
site comparisons can be made only at the level of major anatomical units:
head (horn, cranium, jaws and teeth), thorax (vertebrae, ribs, and sternum),
forelimb (scapula, humerus, radius and ulna), hind limb (pelvis, femur,
patella, tibia and fibula), and extremities (carpals, metacarpals, tarsals,
metatarsals, phalanges, and sesamoids) (p. 168).

The interpretations of element representation based on NISP per element are not as

suitable as those based on MNE for explaining ungulate carcass butchery and transport

patterns. Each specimen is treated as a whole element so that highly fragmented ones,

typically the marrow-filled limb bones, are almost certain to be over-represented. On the

other hand, many studies do not systematically attempt to identify limb shaft portions,

thus limbs will more than likely be under-represented if density-mediated attrition due to

chemical weathering or carnivore activity has impacted the assemblage or if the assemblage

reflects human marrow and grease processing. Since these factors are often not adequately

accounted for, many potentially unwarranted conclusions can be made. For example,

Castaños (1993) in his report on the Magdalenian fauna from Chaves writes,
The elevated degree of fragmentation of the Aragonese sample
produces an over-representation of cephalic elements by a large quantity of
isolated dental pieces that consequently reduces the relative importance of
the long bones represented in good portion by indeterminable diaphysis
fragments. Nevertheless, this circumstance does not totally explain the
limited proportion of ribs and vertebrae, more in the case of the latter, which
presents an elevated degree of conservability in the face of diagenetic factors.
But it is necessary well to suppose that it is an indication of the little frequency
whereupon that part of the animal was transported (Castaños 1993: 16; my

Depending on the method used, this case could be interpreted in many ways. The

frequencies given are percentages of the total NISP. Phalanges and femora are treated as

equal despite the fact that there are 12 times as many phalanges in the ibex skeleton.

However, the admission that indeterminate limb shaft fragments are well-represented

suggests that ibex were intensively processed for marrow and grease onsite. The rabbit

representation at Chaves shows similarities with Picareiro and Suão. The most common

element is the innominate followed by the tibia, mandible, and slightly lower frequencies

of the forelimbs and femur. Ribs and vertebrae are extremely under-represented.

Unfortunately, the majority of the rabbit assemblage was analyzed by another researcher

and the results have yet to be published. Utrilla (1995) reported a preliminary MNI estimate

of 116 rabbits in one level based on innominate frequencies.

The estimation of relative abundance of each species is commonly used to make

statements about their dietary importance. Davidson (1976, 1989) has investigated the

importance of rabbit, red deer and ibex by considering meat weight as discussed previously.

He admits that his main motivation in 1976 was to prove the low significance of the

abundant rabbit remains in Spanish caves (Davidson 1989). Other zooarchaeologists

working in Spain have followed his lead but usually make general statements about the

relative abundance of species based on Davidson’s conclusions. Some have even used

bone weight percentages as a measure of relative abundance (Estévez in Olaria et al. 1985).

A consensus has been reached among scholars that rabbit, while numerically dominant,

was simply too small to have made a significant contribution to the diet. Calculations of

the amount of rabbit meat necessary to equal a single ibex range from 55 to 100 and 88 to

150 for red deer (Davidson 1976, 1989; Villaverde and Martinez Valle 1995). Thus, at

Volcán de Faro where 3,540 rabbit remains were recovered making up 90% of the total

assemblage Davidson (1989) estimated that they only made up 7% of the procured meat.

This reasoning is deeply flawed because Davidson used NISP counts to calculate the

amount of meat. As in all other Spanish cases, each specimen was treated as representing

a whole carcass. Thus, 232 red deer bone specimens were treated as 232 red deer, providing

over 15,000 kg of meat to the cave occupants (Davidson 1989: Table 8.18). Body part tables

show that most of these identified specimens are phalanges, metapodial fragments and

isolated teeth (Davidson 1989: Table 8.10). Femurs were represented by two fragments of

the proximal shaft with the articulation visible. Tibiae by 7 fragments, all either proximal

or distal ends showing the articulation. The forelimb is represented by a proximal scapula

fragment, one proximal and three distal epiphysis fragments (one intact enough to allow

measurement) and one proximal and two distal radius fragments. There is no description

of the rabbit material as it was deemed too insignificant in the face of overwhelming

numbers of red deer. Even if one were to liberally assume each red deer fragment

represented a whole element, at most it could be argued that four animals were evident.

Therefore, only about 265 kg of meat may be represented. Rabbits certainly did not provide

2,655 kg of meat but may have been relatively more important. If the 3,540 remains came

from 100 animals, a reasonable estimate based on NISP/MNI at Picareiro and Suão, then

rabbit may have comprised 25% or more of the meat. This would make rabbit a far more

important resource than many would admit. It would also suggest a great similarity

between the use of Picareiro and Volcán. Because NISP is the basis of counting in Spanish

zooarchaeology, meat weight estimates used to calculate the relative dietary contribution

of rabbit will always greatly exaggerate the ungulate proportion. Since all medium

ungulates appear to be highly and equally fragmented it may be reasonable to use NISP-

based calculations between ungulate taxa but not between large and small game.

In spite of the lack of detailed data on skeletal element representation in the Spanish

sites, there are visible trends in the NISP data that suggest important diachronic shifts in

subsistence in both Mediterranean Spain and Portugal. During the Microlaminar

Epipaleolithic (EMM- Epipaleolítico Microlaminar Mediterráneo 11,000-9,000 bp) of the

SMR the cumulative rabbit NISP falls by more than half to 10,862 while red deer rises to

2,381 and ibex to 1,430 (Table 5.26). Wild boar and chamois increase as well. The

Mediterranean Geometric Epipaleolithic (EGM- Epipaleolítico Geométrico Mediterráneo)

dated 9,000-7,500 has a further drop in rabbit NISP to 1,327. However, while red deer

drops to 379, ibex stays relatively the same at 1,057, suggesting some degree of ibex

specialization at sites such as Tossal de la Roca and Cueva de la Cocina. At Tossal de la

Roca, rabbit exploitation had been stable during the MSM and EMM, but falls dramatically

during the EGM while ibex remained stable throughout the occupation.

Table  : Summary of NISP of fauna from sites in Mediterranean Spain

(adapted from Aura Tortosa & Pérez Ripoll )

Equus sp Bos sp  

Cervus elaphus  ()   ()
Capra pyrenaica 
()   () 
Rupicapra Capreolus   
Sus scrofa 

At Picareiro, the greatest concentration of rabbit bones is in Levels F & G, dated 12,300-

11,700. Rabbit NISP is one third that in all of the Spanish sites for the same time span.

Combined with Suão and Caldeirão, these three sites contain about half the amount of

rabbit from all of the Spanish sites in Table 5.25. Additionally, the number of rabbit remains

drops in Picareiro Level E coincident with the decrease in Mediterranean Spain.

At first glance the decrease in lagomorph remains in Spanish sites appears to coincide

with a greater emphasis on red deer and ibex during the EMM and ibex later in the EGM.

Table 5.26, adapted from Aura and Pérez Ripoll shows the trend in NISP for large taxa in

the SMR from the MSM to the EGM. It appears there is a significant overall shift from red

deer focus to an ibex dominated pattern by the EGM. However, altitudinal and topographic

characteristics determine whether red deer or ibex dominates ungulate assemblages. All

of the EGM sites in their table are located in or near prime ibex habitat and are dominated

by ibex throughout the entire sequence (e.g., Tossal, Cocina, Nerja & Santa Maira). The

disappearance of red deer is due to the absence of occupations from sites on or near the

coastal plain where red deer were exploited during the Magdalenian (e.g., Cova de les

Cendres, Matutano, Volcán & Blaus). So why are there no Geometric Epipaleolithic data

from the coastal plain? The occurrence of marine resources inland and an apparent

mutually exclusive seasonal exploitation of red deer and ibex suggests a coastal-inland

transhumance during the Late Pleistocene as hypothesized for central Portugal. The

absence of red deer could be due to forest expansion on the coastal plain forcing people to

stay on the coast. However, wild boar, roe deer and chamois, typically argued as evidence

for warm, humid Mediterranean forests, decline as well. On the other hand, the decrease

in rabbit appears real since they are ubiquitous and equally abundant in sites dominated

by red deer or ibex. The unique situation of Nerja, near ibex habitat and the Late

Pleistocene/Early Holocene shoreline shows that marine resources were important

throughout the Magdalenian and Epipaleolithic sequence. Aura et al. (1998) note that fish

are five times more abundant than rabbit at Nerja during the Magdalenian (MSM) and ten

times as much during the Epipaleolithic (EMM). A couple of known Geometric

Epipaleolithic sites, now destroyed, were located on the margins of small ponds or coastal

lagoons (Martinez Andreu 2001). It is most likely the EGM settlement was nearer the

coast which was still slightly further out from the present shore. Thus, all of the coastal

sites from this period and earlier are now submerged. A similar pattern emerges during

the Epipaleolithic in Portugal where small shellmiddens (Toledo, Magoito, São Julião,

and Cabeço do Curral Velho) appear on the margins of coastal lagoons.

What are the reasons for the similarities and differences between central Portugal

and Mediterranean Spain? Rabbit exploitation is almost certainly due to its availability

and high productivity although foraging theorists would argue that post-encounter return

rates determine whether or not a resource is exploited (e.g., Stiner et al. 2000). Red deer

are found in small groups and individually for most of the year. Only in the reproductive

season (autumn) do larger groups (male harems) form. Thus red deer was probably taken

through opportunistic encounter hunting for most of the year. Rabbits living in densely

packed warrens were easily “gathered” through setting numerous traps and snares within

their territory and along regular paths (Hockett and Bicho 2000; Hockett and Haws 2002).

Rabbits offered a comparable or better yield of fat and protein from equal amounts of red

deer (Hockett and Bicho 2000)(see also Table 4.9). Why then does rabbit appear to be

more important in Portugal? Were large game resources depressed from excessive hunting

by ever-increasing hunter-gatherer populations? As shown above red deer size in Portugal

did not decrease during the Late Pleistocene (Davis 2002). They were at least as large on

average as those from Mediterranean Spain (both smaller than Cantabrian red deer). One

possibility appears to be the lack of ibex in Portuguese Tardiglacial sites as opposed to

Mediterranean Spain. Current data are probably not sufficient to test this proposition

because of the greater number of sites in eastern Spain as opposed to central Portugal.

The reliance on ibex in Mediterranean Spain is probably due to the fact many sites are

located in ecotones between higher elevations and coastal lowlands. The absence of ibex

in the known Portuguese sites is best explained by the lack of nearby peaks greater than

1,000m. Ibex may not have been very abundant in Late Pleistocene central Portugal due

to the lack of its prime habitat.


Is the decrease in rabbits in central Portugal after 11,000 bp a localized phenomenon

at Picareiro? The other sites in the area with Epipaleolithic occupations such as Bocas,

Papagaio and Pena de Mira have not been analyzed which makes this hypothesis difficult

to test. In fact the collections are too poor to use with confidence. Could the rabbit decline

at Picareiro be due to poor preservation in Levels D and E? This is quite possible given

the low frequency of all taxa. Are there changes in species selection by hunter-gatherers?

Is reduced availability through overharvesting or the level of productivity responsible for

the rabbit decline? Paleoenvironmental changes associated with the Younger Dryas are

difficult to detect in Portugal due the lack of long pollen records and high-resolution micro

and macrofaunal data. If the Younger Dryas impacted Portugal by increased aridity and

cooler temperatures as in northeastern Spain, then preferred rabbit habitat may have been

reduced enough to affect rabbit populations (Hockett and Haws 2002). Many of the Spanish

Mediterranean and Portuguese sites are impacted by erosional events or “ruptures” in

the sequence during the Pleistocene-Holocene transition (Villaverde Bonilla and Aura

Tortosa 1995; Zilhão 1993). At Tossal de la Roca the interior sequence ceased and occupation

shifted to the exterior of the shelter. In sites where rabbit NISP drops, medium ungulate

NISP remains abundant. Does this imply an increase in overall foraging efficiency? The

paucity of sites per millennium might suggest a drop in regional populations thus relaxing

the need for intensive exploitation of “low-ranked” small game. On the other hand, the

first archaeological indications of plant exploitation and significant marine resource use

occurs during this same period. Did the collapse in rabbit populations force hunter-

gatherers into a further broadening of diet to include even lower-ranked plants and

shellfish? While early postglacial red deer size appears smaller on average in Cantabria,

this diminution has been argued as climate-induced (Mariezkurrena and Altuna 1983).

Weinstock (1997) has also noted similar climate-induced reductions in body size for reindeer

in northern Europe. The Portuguese data are insufficient to construct and compare age

profiles to test the hypothesis put forth by Broughton and O’Connell (1999) so the Spanish

data are used here. Age profiles for red deer and ibex in Mediterranean Spain show

continuity in hunting patterns throughout the Paleolithic sequence (Aura et al. 2002). The

differences occur between interior and coastal sites and are likely related to seasonal hunting

strategies and not changes in overall foraging efficiency. This points to the risk involved

in using faunal remains from a few sites to measure foraging efficiency. Differences in

site use and seasonality can lead to erroneous conclusions unless they are accounted for,

which they are commonly not.

Furthermore, as discussed above and in earlier chapters, the intensive use of rabbits

extends back to the earliest Upper Paleolithic. Marine resources are difficult to address

due to coastline changes. Plants were available in fluctuating proportions throughout the

Late Pleistocene but their use is also difficult to evaluate because of the nature of cave and

rockshelter use during this period. Their sporadic occurrence from the Middle Paleolithic

onwards suggests a regional pattern for overall dietary diversity. There does not appear

to be a diachronic trend towards greater dietary diversity or intensified resource use. Diets

were always diverse and carcasses were often intensively processed throughout the Upper

Paleolithic in Mediterranean Spain and Portugal.


Chapter 6: Towards an understanding of Late Pleistocene/ Early Holocene

subsistence and settlement in central Portugal

What can be said of Late Upper Paleolithic and Epipaleolithic subsistence in central

Portugal? If caves and rockshelters and a few small coastal shellmiddens were occupied

for relatively brief periods to process animals, can they be considered representative of

the whole? In most regions, archaeologists consider large and medium ungulates the

basis of subsistence economies because they provide tremendous amounts of energy (kcal)

per unit weight (kg). Small game, fish, shellfish and plants are considered back-up resources

or insurance policies against the risks of stress or failure. According to the Broad Spectrum

Revolution model, evidence of their consumption signals subsistence stress due to

population-resource imbalance. Ultimately, these notions are derived from the focus on

the capture of energy from the environment. The Broad Spectrum Revolution is a

manifestation of the principles underlying optimal foraging theory intended to explain

the apparent global trend in subsistence change toward greater dietary diversity and

resource intensification at the end of the Pleistocene.

In environments where resources are rare or dispersed, animals, and presumably

humans, will be generalized feeders occupying a broad dietary niche. In resource-dense

zones or patchy areas animals and/or humans may be specialized. The Late Pleistocene

of central and northern Europe may have been a resource-dense region where large

gregarious ungulates could be hunted by highly specialized human groups. However,

this may only have been a strictly seasonal specialization. Forest expansion and faunal

turnover to smaller and less dense game at the Pleistocene/Holocene transition forced

people to become more generalized, broadening their diet to maintain energy capture.

On the other hand, increasing population may have played a role in forcing this shift prior

to climate change. In either case, a neat explanation is offered for changes in Late

Pleistocene/ Early Holocene subsistence, and subsequently, settlement patterns.

However, recent ethnoarchaeological studies show that resource rank does not always

correlate with dietary contribution, that men have non-subsistence motivations for large

game hunting, and that overall daily subsistence is often based on the collection of “lower-

ranked” resources by women supplemented by meat (Hawkes et al. 1991, 1997, 2001; Kaplan

and Hill 1992). Furthermore, human nutritional requirements make it extremely difficult

to subsist largely on animals. In regions were animals and plants are available, humans

choose a balance between the two. The broad spectrum diet of Late Pleistocene/Early

Holocene and later hunter-gatherers in temperate latitudes is reflective of the overall

hominid diet that evolved in the last few million years. Current archaeological evidence

shows that the exploitation of plants and small aquatic and terrestrial animals predates

the Pleistocene-Holocene transition in regions where they were available (Kislev et al.

1992; Mason et al. 1994; Richards et al. 2000; Erlandson 2001; Stiner 2001). Any meat-only

diets that may have characterized these latitudes during the Pleistocene are probably highly

specialized adaptations that occurred in times and places of severe climatic conditions

resembling the present-day Arctic and Subarctic. In regions of milder climate this paradigm

should not be considered relevant.

In the Mediterranean region of southern Europe generalized dietary diversity appears


to predate evidence for population increases (seen by numbers of sites per millennium),

although this is debatable (Stiner et al. 2000; Kuhn and Stiner 2001, 2002; Villaverde et al.

1997; Speth and Tchernov 2002; Hockett and Haws 2002). This is especially the case on

the coastal fringe of the Iberian Peninsula excluding northern Spain and possibly Catalunya.

Beyond the Ebro River the archaeological record from the Middle Paleolithic onward is

characterized by the presence of large and medium ungulates, especially red deer and

ibex, but with substantial amounts of small game, mostly rabbit, aquatic resources and

plant remains. The same level of intensity in rabbit processing observed in Magdalenian

sites can be seen in assemblages as early as the Aurignacian at Cova Beneito in

Mediterranean Spain and the Gravettian at Anecrial in central Portugal (Aura et al. 2002;

Hockett and Haws 2002). The Middle Paleolithic record of rabbit use is much less definitive.

These assemblages do not mirror patterns observed in those resulting from carnivores

and raptors nor do they resemble human-created ones. Neanderthals may have no

behavioral analog in modern humans.

Recently, Hockett and Haws (2002) reviewed current models explaining rabbit

exploitation in the western Mediterranean. In the first one, Villaverde et al. (1997) argued

that increased rabbit hunting in the Upper Paleolithic indicates a reduction in mobility.

Because rabbits are a dense territorial resource Early Upper Paleolithic people became

tethered to them. This was further supported by the nature of medium ungulate prey

behavioral ecology. Rabbits were a ubiquitous resource that served as an insurance policy

while small foraging groups moved seasonally between the coastal plain where they

targeted red deer and the interior where they hunted ibex.

In the second model, Stiner et al. (2000) proposed an alternative for overall small

game exploitation based on population pressure on resources (see chapter 2). According

to their model the increased role of lagomorphs signals depression of higher-ranked

resources due to over-harvesting. Therefore, the intensive utilization of rabbits in the

Upper Paleolithic of Iberia would suggest an earlier occurrence of the Broad Spectrum

Revolution, as modified to acknowledge aquatic resource use prior to the Pleistocene-

Holocene transition (Stiner 2001; Stiner and Kuhn 2001, 2002). The Stiner et al. (2000)

model primarily focuses on hares rather than rabbits. As discussed earlier, hares differ

from rabbits in many important ways. Hares are solitary, live in nests on the ground and

though capable of rapid bursts of speed to escape predation will freeze in place as an

initial defensive posture. Keen hunters may be able to stalk, approach and dispatch hares

from a relatively short distance. Alternatively, drives could be used in certain cases. Rabbits

on the other hand are easily gathered in large quantities through methods requiring little

effort. Despite their overall speed, they may not fall into a general “quick-moving” category

put forth by Stiner et al. (2000).

An alternative model for rabbit hunting and overall small game exploitation was

developed by Hockett and Haws (2002). This model invokes prey behavior and climate to

explain the temporal trends in rabbit hunting in the western Mediterranean. Rabbits are

an attractive resource for humans not just because they are ubiquitous and abundant but

because they are relatively easy to procure in large numbers. In other words, rabbits may

have a much higher return rate than simple meat-weight and caloric energy models suggest.

In addition, as Hockett and Bicho (2000) noted, rabbits offer a better nutritional package

of protein and fat than deer, despite the low-fat, ‘rabbit starvation’ claims for other regions

(e.g., Harris 1987; Speth and Spielmann 1983).

The models adopted by archaeologists working in Iberia all focus on medium and

large ungulates as the basis of subsistence with settlements arranged in order to procure

these resources. The data may not be sufficient at the present to test the idea that ungulate

exploitation was highly specialized in central Portugal as it may have been in Mediterranean

Spain. Larger assemblages in that region have allowed the construction of mortality profiles

that show the regular hunting of animals ~3 years old (Aura et al. 2002). This pattern

varies between the coast and interior and probably has more to do with seasonal differences

in herd structure than dietary focus. The assemblages from the Portuguese sites are not

large enough to construct useful mortality profiles. Coastal Magdalenian sites are unknown

and the few dated to the Early Holocene are shellmiddens whose terrestrial game

component was either non-existent or has not been studied. Analyses of Picareiro and

Suão show differences in large game utilization. At Picareiro, medium ungulate carcasses

were brought to the cave and intensively processed for maximum nutritional yield. Suão

was used for this purpose to a much less degree. If large game was processed in the site,

very little was consumed onsite. In both sites, rabbits were procured and processed in

large numbers throughout the Late Pleistocene sequence.

If medium ungulates were heavily exploited the impact was probably not severe

enough to “depress” populations. In Spain, where red deer specialization is said to occur

as well, the mortality data show no significant changes through time. The biometric data

from Picareiro and Caldeirão show no size diminution in red deer, arguably the most

exploited ungulate. This does not agree with the prediction of the diet breadth model that

low-ranked resources will not be added to the diet until higher-ranked resources offer

diminished returns to the point at which it becomes profitable to add new items. The only

way to satisfy the model is to suggest that rabbits were not in fact low-ranked resources

despite the discrepancies in meat weight and caloric value. This explanation implies that

the commonly held view that large game outrank small game is seriously flawed.

Given that the interior caves and rockshelters do not represent all types of settlement

what reason is there to assume that the faunal remains found in them represent the full

repertoire of diet and subsistence? Comparative studies of these sites in Portugal and

Mediterranean Spain suggest they are primarily specialized carcass processing locations

rather than residential camps. Milder climate probably meant that much of the terrestrial

game processing and consumption took place in open air sites where organic preservation

is extremely poor due to the acidic soil conditions. Plant and aquatic resource utilization

was likely much higher despite their low frequency in caves and rockshelters. Plant

gathering and fishing took place in areas where they were locally abundant. The same

can be said for terrestrial game exploitation. The name of the village nearest Picareiro is

Covão de Coelho, meaning ‘rabbit cave.’ This would suggest that rabbits have been very

abundant for a long, long time in the area.

The present landscape near Picareiro and Suão has been altered by thousands of years

of human forest clearance, agriculture, silviculture and domestic animal grazing. The

limestone uplands probably had more extensive deciduous and evergreen Mediterranean

forests than the maquis or garrigue that characterizes the natural vegetation today. We

can only speculate on past plant availability. The potential for plant resource use discussed

in chapter 4, while speculative, should not be dismissed simply because plant remains are

absent in specialized animal processing sites.

How significant were marine resources to the Late Pleistocene/Early Holocene diet

in central Portugal? The archaeological evidence from this period shows that people not

only utilized coastal resources near the shore but transported them considerable distances

inland. Were people moving residence seasonally to the coast to exploit marine resources,

or were they making special trips to the coast to collect food and other resources to bring

back to interior residences? Ethnographic observations of groups in central California

record both patterns. The Coast Yuki moved residence seasonally within a 25 mile area

between the coast and inland coastal range (Gifford 1939). On the other hand, the Yokuts

were observed to travel at least 50 miles to collect molluscs for transport inland (Pilling,

1950). Long distance transport of shells inland has also been observed in New Zealand

(Coutts and Higham, 1971), Tasmania (Coon 1971) and Tierra del Fuego (Massone 1987).

In the Santa Lucia Mountains of central California, substantial prehistoric shellmiddens

comprised of marine mussels are found 25-30 km inland (Jones and Richman 1995). In

Peru, early Holocene hunter-gatherers transported shellfish 100 km inland (Engel 1973).

Foraging models suggest that fulfillment of energy needs may not have been the motivation

for long-distance shellfish transport (Jones and Richman 1995).

For the past several decades, archaeologists have been divided on the nature and

timing of coastal adaptations. Coastal adaptations were argued to have appeared as a

result of population growth and the stabilization of sea levels in the early Holocene (Binford

1968; Cohen 1977; Osborn 1977; Yesner 1980, 1987). The reasons given to explain why

coasts were not exploited appreciably during the Pleistocene are:

1) human populations did not reach the point necessary to broaden the diet
to include low-ranked shellfish and marine mammals,
2) sea levels were fluctuating enough to preclude the formation of rich
productive estuaries
3) that overall ocean productivity was lower because of lower global CO2,
4) continental shelves were exposed which left very deep unproductive water
off the coast (Bailey and Parkington 1988).

Erlandson (2001), in a thorough review of the worldwide evidence for coastal resource

use, shows quite demonstrably that coastal adaptation has a long history albeit in restricted

areas. The primary reason for the lack of coastal sites is the fact that sea level has inundated

or destroyed most Pleistocene coastal sites. The sole factor in the presence of coastal sites

from this period is that areas of steep bathymetry enabled site visibility (Erlandson 2001).

In these areas, the distance from the present shoreline to the glacial one is only a few

kilometers because of the narrow continental shelf. Additionally, sites located above the

Last Interglacial shore could be preserved because sea level has not reached that height

and inundated them. This explains why coastal resources are common in Upper Paleolithic

sites in Cantabria and OIS 5e sites are preserved in North Africa. Since evidence for coastal

exploitation exists in areas where it could be preserved, why assume the lack of evidence

in other coastal areas means that it never existed in the first place?

Despite the seemingly obvious fact that Pleistocene coastal sites are underwater many

refuse to acknowledge that Paleolithic people utilized the coast very extensively because

they view marine resources as inferior to terrestrial ones (Osborn 1977; Yesner 1980, 1987;

Bailey 1978). This stems from the use of energy-based economic models like site catchment

analysis and optimal foraging theory. The most extreme view was promoted by Osborn

(1977) and Bailey (1978). Both considered the protein and energy value of shellfish

inadequate to meet human dietary needs. Hence, Bailey estimated that 150,000 cockles

would be needed to equal a single red deer. Therefore, shellfish could only serve as a

marginal or starvation resource in stressful times.

Akazawa (1988) raised important points in relation to dietary (caloric) contribution

and relative importance of shellfish. Using seasonality considerations made by Koike, he

reconsidered the data from Isarago in Japan. In a previous study of the shellmidden,

Suzuki estimated the dietary contribution of shellfish based on calculation of the volume

of shells in the midden and the weight of the animals. Assuming a year round occupation

for 25 years by 30 people, the inhabitants would have consumed 19g of protein and 134

kcal/day from shellfish. This suggests that shellfish were of minor importance to the diet

despite their numerical dominance. Akazawa argued that if the site was only occupied

for three months, then 48g of protein and 265 kcal/day were obtained from shellfish.

Thus, during certain times of the year, especially early spring and summer in this case,

shellfish were of considerable importance given the large quantities collected in a short


A similar conclusion was reached by Meehan (1982) in her ethnographic study of the

Gidjingali of Australia. For them, shellfish are a critical resource during the summer wet

season. Though shellfish were not a critical source of energy, only amounting to 6-17% of

calories during the year, the Anbarra community gathered shellfish as often or more than

any other food. More significant than the caloric yield was perhaps the ease in collecting

with minimal processing that allowed women and children to provide a stable, reliable

resource. As with many modern hunter-gatherers, the Anbarra purchase about half of

their food, mainly flour, sugar and rice. This apparently has not affected the amount of

animal flesh consumed but has taken the place of tubers, nuts and fruits in the overall diet

(Meehan 1982).

Erlandson (1988) has shown that many shellfish are not poor protein sources (see

Table 4.14). Regardless of the amount, the protein quality of shellfish is the highest of any

source for humans (Wing and Brown 1979). The protein proportions in shellfish are best

suited for human metabolism. Furthermore, given the risks involved in a high protein

diet, it would seem unrealistic to rank food items according to the amount of protein.

Generally, the occurrence of marine fish and shellfish in coastal middens and caves/

rockshelters in the interior is thought to represent the initial use of these resources by

human groups under stress from population/resource imbalance. Indeed, energy-based

foraging models assume these resources rank lower than terrestrial ones and would

therefore only be used when higher ranked resources were depressed. Many studies over

the last decades have concluded that shellfish could never have been very important dietary

components due to the low meat weight relative to larger terrestrial game (Osborn 1977;

Bailey 1978; Yesner 1980, 1987; Perlman 1980). Again, Bailey’s (1978) assertion that over

150,000 cockles would be needed to equal the meat weight of a single red deer illustrates

this line of thought. Subsequent researchers have concluded the same, except Erlandson

(2001). Shellfish therefore, are thought to represent a post-Pleistocene adaptation in the


face of ever-increasing demographic pressures, whether it is population growth or

“packing.” This would apply to ocean fish and marine mammals since their appearance

would be seen as further evidence of increased diversity.

An alternative explanation put forth by others is that coastal resource availability

was low throughout the glacial period because continental shelves were too steep to permit

productive fisheries and the broad estuaries and lagoons necessary to support large colonies

of shellfish (Schubel and Hirschberg 1978). Only after sea level rose in the Late Pleistocene

culminating with the Atlantic transgression were these marine conditions met. Therefore,

the addition of marine foods is due to superabundance. This, of course, would only fit the

foraging theory models if these resources were ranked higher than the ones already

included in the diet. If they do not rank higher then population pressure again would be

used to explain their inclusion.

However, the evidence suggests shellfish have long been a food source for humans,

even during the Last Glacial Maximum in Cantabria (Clark 1983; Clark and Straus 1986;

Erlandson 2001). The main obstacles in recognizing that coastal exploitation was not merely

a localized and rare phenomenon during the Pleistocene are twofold. The first is the

submergence of the Pleistocene shoreline at the end of the Last Glacial. We simply lack a

substantial amount of prehistoric human territory. There is no reason to believe the unique

Tejo estuary and others in Eurasia were indicative of a change in the overall productivity

of coastal regions globally. Estuaries, lagoons and marshes would have formed on exposed

flat lands of the exposed continental shelf during lowstands (Dias et al. 2000). The geologic

record of coastal landforms was mostly destroyed and remnants are preserved underwater.

El Juyo
La Riera
Altamira Rascaño


Atlantic Ocean

Cova Fosca
Buraca Grande
Lapa do Picareiro Caldeirão Cova dels Blaus
Bocas Cueva de la Cocina
Lapa do Suão
Volcán de Faro
Les Mallaetes
Parpalló Santa Maira
Tossal de la Roca
Cova de les Cendres

Vale Boi

Nerja Mediterranean Sea


Figure  : Map of Iberia showing the main sites and their position
in relation to the coastline during the Late Magdalenian

The absence of a geologic and archaeological record of coastal resource abundance is not

evidence of its absence.

The assumption that rich marine and estuarine biota could only form after sea levels

reached their current position and “stabilized” now for several millennia is weak at best.

The immediate response by Mesolithic foragers to the “sudden” stability and productivity

of the coast shows that these biological communities did not require millennia to form.

They were already present but further out, living in now submerged estuaries, lagoons

and marshes. To argue otherwise, as many have repeatedly done, is to abandon


Table : comparison of shellfish and large mammal

energetic values and return rates
kcal/g return rate



red deer  
wild boar 

The second obstacle is the use of energy as the sole currency in economic models.

Table 6.1 shows the energetic values and estimated return rates of shellfish compared to

terrestrial game. The conclusions drawn from such comparisons by Osborn (1977), Bailey

(1978), Yesner (1980, 1987), etc. seem self evident. However, as Tables 4.12 and 4.14 showed,

it is clear that some shellfish types rival terrestrial animals in protein and fat content.

They also offer carbohydrates which are completely lacking in terrestrial mammal meat,

making shellfish an important resource in times of low plant availability. The fat content

of fish and shellfish peaks during summer as they exploit the abundant food resources

created by the upwelling of nutrients (Table 6.2). This would have been the optimum

time of the year to utilize coastal resources as plant availability would be lower and

terrestrial game fat reserves depleted due to the summer drought. Furthermore, as Perlman

(1980) suggested, shellfish might outrank deer because of the lower risk of failure in shellfish


Table  : Seasonal nutritional composition of sardines

Sardine Summer/Fall Winter Spring


Data from Tabela da Composição dos Alimentos Portugueses

(Gonçalves Ferriera & da Silva Graça  )
All analyses conducted on fresh fish

Based on these data, the coast may have been an attractive place to live during much

of the Late Pleistocene. Recent oceanographic work shows the LGM coastline was situated

30-40 km away in most of Estremadura (Dias et al. 1997, 2000). This is a critical problem in

evaluating the nature and timing of marine resource use because the evidence is 100-150

m underwater. In fact, it is not until the end of the Pleistocene that we see the first coastal

shellmiddens in Portugal, when sea level was close to its current level. Therefore, it is

tempting to suggest that marine resource use at the end of the Pleistocene represents a

new adaptation. The evidence of seals and limpets from the coastal Mousterian site,

Figueira Brava, dolphin from the interior Gravettian site, Lagar Velho, and limpets from

Val Boi, a Gravettian and Solutrean site in Algarve, should be enough to show this is a

Buraca Escura
Buraca Grande
N Lagar Velho


Atlantic Ocean Picareiro
Carneira Passal Casal do Cepo
Vale Comprido
Casa da Moura
Lapa do Suão
Porto Dinheiro
Lapa da Rainha
Vale Almoinha

Rua de Campolide

Poço Velho

Vale Boi

Figure : The coastline of Portugal during the Last Glacial Maximum

fallacy. One could also point to Cantabria, where the LGM coast was only 7-9 km away

and there is solid evidence for shellfish exploitation during the Solutrean. Substantial

amounts of shellfish were found in the Solutrean levels at Parpalló (Pericot 1942). Cockles

and mussels were recovered in Solutrean contexts at Nerja (Jordá 1986). These sites show

that the coast was settled and exploited prior to the end of the Pleistocene and regardless

of climatic conditions.

Though evidence for Solutrean marine food exploitation in Portugal is non-existent

some hypotheses can be made using the geological record. In general, coastal zones are

highly productive where upwelling occurs. These areas are often the focus of human

subsistence (Perlman 1980; Bailey and Parkington 1988). Today, a fairly strong summer

upwelling of cold, deep, nutrient-rich waters occurs off the western Portuguese coast.

This is driven by the Trade Winds and northward flow of the Canary Current along the

northwest African coast (Abrantes 2000). Though not as strong as the one occurring off

the coast of Peru, (60-90 g C m-2 yr-1 off Portugal; 345 g C m-2 yr-1 off Peru) upwelling along

the western Iberian margin provides sufficient nutrients to make the Portuguese coast a

highly productive marine resource zone (Fiuza 1983; Abrantes 1988). Using diatom

abundance in deep sea cores off the coast north of Estremadura, Abrantes (1988, 1991)

noted fluctuations in upwelling intensity over the last 100,000 years. During OIS Stage 3

upwelling intensity was roughly equal to the present. However, paleoproductivity

increased during Stage 2 culminating in the LGM. Upwelling intensity was an order of

magnitude greater than the present making the coastal zone more fertile than today. This

was probably due to intensification of the Trade Winds (Abrantes 2000). During the

deglaciation, around 15-12 kya, upwelling intensity decreased substantially but was still

3-7 times stronger than today. Interestingly, the Pleistocene/ Holocene boundary marks

a time when the coastal upwelling was at its weakest in the last 20,000 years. By the Early

Holocene it dropped below current levels only to rebound in the last few thousand years.

Independent studies on organic and inorganic carbon plus C37 alkenones from pytoplankton

in marine sediments by Pailler and Bard (2002) confirm the intensity of the upwelling

during cooler periods and its reduction during warm ones. Using a combination of marine

proxy indicators, CaCO3, barium and diatoms, Thomson et al. (2000) found that the

enhanced productivity of the ocean off the Portuguese coast during cold climatic events

peaked at the end of glacial periods when sea levels rise.

Given these facts it is likely that the western coast of Portugal during the Solutrean

was an extremely attractive place for human settlement and subsistence. Marine resources

would have been abundant but the terrestrial resources of the coast would have been

relatively poor. In areas of the world where coastal upwelling occurs, the adjacent lands

are semi-arid to arid (DiCastri 1981). This is true for the coast of Portugal, northwest

Africa, the skeleton coast of Namibia, the coastal deserts of Peru and the coast of California.

The region with the most intensive upwelling, Peru, also has the greatest coastal aridity.

The patterns of trade winds, cold ocean currents and coastal geography create this situation.

It could be argued that upwelling intensity is correlated with coastal aridity (Shi et al.

2000). Recent studies of the Benguela upwelling system off Namibia show a correlation

between changing upwelling intensity and terrestrial vegetation.

For central Portugal, pollen cores and charcoal studies show that vegetation at higher

elevations was dominated by cold, arid Artemisia steppe. The land area now submerged

is thought to have been mainly sandy dunes with little vegetation (Daveau 1980; Zilhão

1997). Most of the research on the continental shelf has focused on the northern Portuguese

coast and many of the observations for the Estremaduran coast are based on extrapolation

(Dias et al. 2000). During the LGM, the continental shelf was much steeper and wave

activity would have been strong. Dias et al. (2000) argued that most of the sediment from

terrestrial river systems flowed through the deep submarine canyons because the slopes

between them are largely rocky and devoid of recent sediment. Sedimentation was high

because lower sea levels caused lower base levels for streams. In addition, Daveau (1980)

suggested greater annual rainfall and spring ice and snow melt in the mountains added

even more sediment to streams. While sea levels may not have been at their maximum

low long enough for soil development and forest colonization, they were considerably

lower, up to 100 m, prior to and after the LGM for thousands of years. This stability is

seen in the preservation of relict abrasion platforms, sea cliff remnants and off shore bars

on the shelf (Dias et al. 2000).

Much of the now-submerged land may have been moderately forested during the

Gravettian and Magdalenian. With the productive Mediterranean plants and certain types

of game, especially rabbit and wild boar, restricted to unknown refugia during the LGM,

ibex and red deer would have been the main large game in the uplands, with horse in

grassland zones. In order to maintain dietary balance in the face of reduced terrestrial

plant productivity and rabbit populations, humans would have relied heavily on coastal

resources, including fish, shellfish and marine mammals, especially during summer.

Because the coast was much farther away, there is little evidence for inland transport of

marine resources 25-40 km as there is after 10,500 bp. The majority of sites with preserved

faunal remains would have been 30-60 km inland while those from the Gravettian and

Magdalenian would have been 40-50 km inland. Where the bathymetry is steeper, sites

with faunal preservation that would have been near the coast during lowstands contain

evidence of marine resources transported inland. Vale Boi, in Algarve, illustrates the

point as it contains numerous limpet shells, yet it was at least 20 km inland during its

occupation. The Gravettian site of Lagar Velho, with dolphin remains, would have been

15-20 km inland. Picareiro would have been almost 40 km inland when fish and shellfish

were brought to the site during the Magdalenian. Even Lapa do Suão would have been 25

km inland and it contain marine fish and shellfish. Thus, the perception that this behavior

reflects a new subsistence strategy or a diversification of the diet is illusory.

The Termination IA/Dryas I/Heinrich I cold period would have presented serious

problems for hunter-gatherers. Sea surface temperatures were much cooler than during

the LGM. Upwelling intensity was still strong but not as high as the LGM. The dry

summer/ wet winter pattern was replaced by a dry summer/ dry winter one. Pollen

cores indicate a further decrease in arboreal pollen. Perhaps not surprisingly, the

archaeological record for this period is extremely poor. In fact, there appears to be a real

hiatus in radiocarbon dates between 14,500 and 12,500 bp. Whether or not this represents

a serious decline in regional population remains to be tested. People would have faced

year-round drought stress. Terrestrial game was probably less available and in poorer

condition. Coastal resources, especially shellfish with its minimal carbohydrate, would

have been even more necessary. Fish and marine mammals may have been important

sources of protein and fat. People may have shifted settlement locations to the coast year

round. These sites would all be underwater now if they existed.

When temperatures increased again after 13,000 bp, corresponding to the Bølling/

Allerød phase, arboreal pollen increased dramatically. This period appears to be as warm

as today with perhaps more humidity. Mixed evergreen and deciduous Mediterranean

vegetation spread along with rabbit and wild boar, perhaps to the detriment of ibex and

chamois. With plant productivity high, the Magdalenian foragers would have had a much

higher diversity of available resources by 12,500 bp. Plant carbohydrates would have

eased the risks inherent in high protein diets and allowed for more balanced nutritional

diets. Combined with the still productive ocean, these groups may have been much

healthier and able to grow population. The increased number of sites dated to this period

offers a striking contrast to the Early and Middle Magdalenian.

Ironically, the widespread appearance of marine resources at the end of the Pleistocene

coincides with a decrease in upwelling intensity and marine productivity. This does not

mean that the coast was no longer an attractive zone. It was at least as productive as

today. However, the terrestrial environment rebounded with higher plant productivity

and increased rabbit populations. Interior sites were once again full of rabbit bones whereas

they were greatly reduced during the LGM. Thus, people had access to rich marine and

terrestrial biomes. The dry summer/ wet winter pattern meant that the most stressful

period of the year was likely summer. Terrestrial game would have lower fat reserves.

Waterfowl would have migrated north to temperate latitudes. Underground plant storage

organs such as bulbs, roots and tubers may have been the best available plant resources.

The best food source would have been fish and shellfish. Their fat reserves would be high

as this part of the year is the growing and reproductive season.

Human settlement would have been focused on the coast. By early fall, when rainfall

returned, nut-producing trees and game animals would be the prime resources. Settlement

could have shifted back to the interior. With the rains, the polje near Picareiro and other

sites in the Serra d’Aire would have filled with water, possibly for longer periods than

present. Freshwater crustaceans would have attracted waterfowl, game and humans alike.

During the mild winter, game animals and stored nuts would have formed the basis of a

nutritionally balanced diet. By early spring, rains again would bring lush vegetation

providing ample supplies of greens and better provisioned game.

This adaptation continued in the Epipaleolithic after 10,000 BP when we see evidence

of shellfish exploitation in small coastal estuaries at sites such as São Julião, Magoito Curral

Velho and Toledo. The apparent settlement shift to include the coast is simply an artifact

of sea level changes. The transportation of shells inland to Bocas, Casal Papagaio and

Pena de Mira is due to the shortening of the distance to the shore. As noted above, there

is no reason to suggest that transporting coastal resources inland was a novel characteristic

of the Epipaleolithic.

The marked shift in settlement from Estremadura to the Tejo basin at the beginning

of the Mesolithic occurred when the Atlantic Transgression created a huge productive

estuary. This near-complete abandonment of Estremadura came at a time when coastal

upwelling intensity was at its lowest in 200,000 years (Abrantes 2000). The shutdown of

this system led to a collapse of the subsistence and settlement system that had been in

place for millennia. Coastal resource availability may have decreased but the transgression

enabled people to reorganize their settlement to focus on the rich and very large Tejo

estuary. This would explain the abandonment of Estremadura and the sudden occurrence

of sites along the margins of the Tejo estuary and small tributaries flowing into it. The

terrestrial resources found in these sites were the same as those used before. There does

appear to be greater aquatic resource diversity but this probably reflects site visibility and

the nature of the estuarine ecosystem, not diet breadth expansion due to population

pressure. However, the concentration of sites may have created local densities not possible

during earlier periods.

Unfortunately, no taphonomic studies have yet been made on the faunal assemblages

from the large Muge shellmiddens. The appearance of cemeteries and structures at Moita

do Sebastião suggests a degree of permanence if not sedentism. The lack of any comparable

open sites during the Upper Paleolithic and Epipaleolithic is further evidence that social

and settlement organization was quite different.


The human adaptations to changing Late Pleistocene climate and environments were

not the result of long-term positive demographic trends requiring a broadened diet niche.

Rather, they appear to have been geared towards diversity from the Early Upper Paleolithic

onwards (Hockett and Haws 2002). Resource intensification is equally apparent throughout

the Upper Paleolithic as rabbit exploitation patterns are similar from the Gravettian to the

Final Upper Paleolithic in both Portugal and Spain. Specialized sites for processing rabbits

and medium ungulates seem to be the only type of faunal-bearing site represented in the

limestone massif. These are all caves and rockshelters. However, there are still few sites

with enough faunal remains to test this hypothesis. The open-air sites seem to represent a

range of sites but almost none are large enough to be considered long-term residences.

The lithic technological studies of settlement patterns indicates a fairly high degree of

residential mobility though not to the degree seen in northern Europe (Thacker 2000).

Seasonal movements between the coast and interior allowed hunter-gatherers to maintain

dietary balance throughout the year. Similar adaptations are apparent in Mediterranean

Spain. The different resource use patterns in Cantabria are due to differences in climate,

geography and environment.

At present, the data from Portugal indicate that climate and paleoenvironmental

conditions best explain the human adaptations in the Late Pleistocene and Early Holocene.

Evidence for increased population size or density is not apparent. The rise in numbers of

sites and occupied area is an artifact of rising sea level. Diets appear to have been equally

diverse throughout the Upper Paleolithic. Specialized hunting of rabbit and ungulates

corresponds to their natural abundance on the landscape. In addition, there is no evidence

of resource depression forcing people to adopt low-ranked resources.

The ‘Tardiglacial paradigm’ needs to be thought of not as a culmination of long-term

progressive trends towards a broad spectrum adaptation but rather as a situational model

that characterizes localized adaptations to changing environments by hunter-gatherers.

The unique site of Nerja gives the false impression that people only began exploiting a

wider variety of coastal resources in the Upper Magdalenian. The position of the coast is

important. The cave was always relatively close to the sea, even during the glacial

maximum. However, the central place foraging models illustrate how only a few kilometers

can make a difference in transport decisions (Bettinger et al. 1997; Bird and Bliege Bird

1997, 2000). There is also the assumption that the cave functioned as a residential camp

during the Magdalenian. Changing site function will be reflected in the kinds of resources

that will be brought to the site and subsequently deposited.

A serious problem in tracking increased resource utilization through time is the use

of NISP. Increases in bird NISP of the amount cited by Villaverde and Martínez Valle

(1995) and Aura and Pérez Ripoll (1992) could mean that only one bird was killed in each

level but more bones survived in the more recent level, thus creating the illusion of greater

use of birds.

Population levels are difficult to assess. There are very few sites and those that exist

are repeated short-term occupations by small groups. There are few if any large open-air

base camps in either central Portugal of Mediterranean Spain. Both regions appear to

have relatively low population densities and a moderate degree of residential mobility

even in times of “intensification” and “diversification.” Straus et al. (2000) show site

distributions for each cultural phase of the Upper Paleolithic in Iberia. The few number of

Aurignacian and Gravettian sites indicate low populations during the Early Upper

Paleolithic. Site numbers increased substantially during the Solutrean. However, there is

a decrease in the Magdalenian just at the time of increased dietary diversity and resource

intensification argued by proponents of the ‘Tardiglacial paradigm.’

The models proposed by Zilhão and Bicho cannot be fully tested without more data

from other cave sites but open-air contexts as well. The pattern observed at Picareiro and

Suão suggests a specialized function of caves and rockshelters as carcass processing stations

not residences. Therefore, without subsistence information from the open-air sites, the

overall subsistence and by implication, settlement pattern cannot be determined. With

chert ubiquitous across the landscape, it is unlikely that settlement patterns will be

necessarily due solely to lithic technological organization. Subsistence practices, including

all of the economic and nutritional decisions made by Upper Paleolithic people, must

have played a dominant role in settlement location. Here the Bicho model can be enhanced

by applying the results of this dissertation.

Some directions for future research

There are several lines of research that need to explored to fully test the ideas presented

above. First, more nutritional data on edible wild plants and animals need to be

systematically collected. Although utility indices derived from experimentation may not

accurately measure economic and nutritional utility they could provide a comparable data

set to the Great Basin studies. Body part utility indices for wild boar, red deer, roe deer,

ibex and chamois would permit a better understanding of skeletal element representation

in Upper Paleolithic sites in Portugal. Nutritional data on the full range of available fish

and shellfish are also needed.

Future fieldwork should involve systematic surveys along the present coast of

Estremadura. Near Nazaré, coastal sand dunes contain open-air sites from the Late

Pleistocene and Early Holocene. There are also numerous caves and rockshelters in the

limestone areas between the Serras de Aire and Candeeiros that remain to be explored

archaeologically. At Picareiro there are undated levels that extend back to the Middle

Paleolithic that need to be opened up in order to investigate problems such as the nature

and timing of rabbit hunting in central Portugal. This has important implications for

current models (Hockett and Haws 2002).

Due to the high-energy waves that shape the western Portuguese coast, the likelihood

that underwater open-air sites could be found is very slim. Survey and exploration of

underwater caves may be a better avenue to pursue. During the Late Pleistocene, when

rivers such as the Tejo were downcutting, people likely camped near their banks. These

locations are almost unknown for the Pleistocene with a couple of notable exceptions.

Because the Atlantic transgression would have rapidly inundated the lower reaches of the

Tejo through low energy processes, the likelihood that Late Pleistocene sites are buried

under several meters of recent alluvium is probably high. Deep coring of these sediments

may be a worthwhile endeavor.

The taphonomic analyses of Picareiro and Suão highlight the need for additional sites

to be studied in the manner. Excavation methodology is generally quite good at present.

Techniques are adequate for recovering the smallest bone specimens and most recognize

the need for keeping all of the remains. However, the recovery of plant remains is not

being systematically undertaken. Flotation needs to be done but so far all plant parts are

recovered only through wet-sieving sediment samples. A new project in Estremadura is

identifying plant exploitation through use-wear and starch grain identification. This holds

promise for recovering evidence of plant use in the open-air sites as well as caves and



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