2010
Table of Contents
ACKNOWLEDGEMENTS ................................................................................................................................................... 5
CHAPTER 1 INTRODUCTION AND SCOPE OF WORK .............................................................................................. 6
1.1 TRADE AND MIGRATION: THE ARCHAEOLOGICAL VIABILITY................................................................................... 7
1.2 THE ARCHAEOLOGICAL SIGNIFICANCE OF INSECTS................................................................................................... 9
1.3 AIMS OF THE STUDY ................................................................................................................................................... 11
1.4 APPROACHING A METHODOLOGY.............................................................................................................................. 11
1.5 STRUCTURE ................................................................................................................................................................. 13
CHAPTER 2 MORPHOLOGICAL AND ECOLOGICAL CONCEPTS ....................................................................... 14
2.1 INTRODUCTION ........................................................................................................................................................... 15
2.2 MORPHOLOGY ............................................................................................................................................................ 15
2.2.1 The Cuticle .......................................................................................................................... 16
2.2.2 The Chitin ........................................................................................................................... 18
2.2.3 The Head ............................................................................................................................. 21
2.2.4 The Thorax .......................................................................................................................... 24
2.2.5 The Abdomen ...................................................................................................................... 25
2.3 IDENTIFICATION METHODS ........................................................................................................................................ 26
2.3.1 Modern Insects .................................................................................................................... 26
2.3.2 Fossil Insects ....................................................................................................................... 28
2.4 ECOLOGY .................................................................................................................................................................... 30
2.4.1 Components ......................................................................................................................... 31
2.4.2 Synecology ........................................................................................................................... 33
2.4.3 Ecological Constraints ........................................................................................................ 36
2.4.4 The Ecology of Invasive Species......................................................................................... 38
CHAPTER 3 METHODOLOGICAL REVIEW ............................................................................................................... 40
3.1 INTRODUCTION ........................................................................................................................................................... 41
3.2 DATA COLLECTION..................................................................................................................................................... 41
3.3 PROCESSING METHODS .............................................................................................................................................. 41
3.4 PALAEOECOLOGY ....................................................................................................................................................... 42
3.4.1 Theoretical Perspective ....................................................................................................... 42
3.4.2 Background to the Methodology ........................................................................................ 42
3.4.3 Methodology ........................................................................................................................ 44
3.5 BIOGEOGRAPHY .......................................................................................................................................................... 44
3.5.1 A Brief Literature Review ................................................................................................... 44
3.5.2 Approaching the Problem ................................................................................................... 45
3.6 ISOTOPIC ANALYSES ................................................................................................................................................... 46
3.6.1 History ................................................................................................................................. 46
3.6.2 Theoretical Basis ................................................................................................................. 47
3.6.3 Methodology and Objectives ............................................................................................... 49
3.7 PHYLOGEOGRAPHY .................................................................................................................................................... 50
3.7.1 The History of Ancient DNA .............................................................................................. 50
3.7.2 The Extraction of Ancient DNA from Samples ................................................................. 51
3.7.4 Ancient DNA Authentication Criteria ................................................................................ 53
3.7.5 Targeting Mitochondrial DNA: The Aims of the Project .................................................. 53
CHAPTER 4 THE PALAEOECOLOGICAL APPROACH: .......................................................................................... 55
4.1 INTRODUCTION ........................................................................................................................................................... 56
4.2 CASE STUDY 1: 7-15 SPURRIERGATE, YORK (SITE CODE: 2000:584) ....................................................................... 57
1
4.2.1 Introduction ........................................................................................................................ 57
4.2.2 Processing Methods ............................................................................................................ 57
4.2.3 Results and Analysis of the Data ........................................................................................ 58
4.2.4 Palaeoenvironmental Reconstruction of Roman Spurriergate ......................................... 69
4.2.5 Palaeoclimatic Reconstruction ........................................................................................... 73
4.2.6 Discussion: The Environment and Climate of 7-15 Spurriergate and Implications for Culture Contact 74
4.2.7 Summary ............................................................................................................................. 76
4.3 CASE STUDY: 16-22 COPPERGATE, YORK PERIOD 4B ............................................................................................... 76
4.3.1 Introduction ........................................................................................................................ 76
4.3.2 Background and Processing Methods ................................................................................ 76
4.3.3 Results and Analysis of Data .............................................................................................. 78
4.3.4 Palaeoenvironmental Reconstruction of Anglo-Scandinavian Coppergate ..................... 98
4.3.5 Palaeoclimatic Reconstruction ......................................................................................... 103
4.3.6 Discussion: The Environment and Climate of Period 4b 16-22 Coppergate and Implications for Culture
Contact ....................................................................................................................................... 104
4.3.7 Summary ........................................................................................................................... 109
4.4 CONCLUSION ............................................................................................................................................................. 109
CHAPTER 5 GRAIN PESTS: AN ARCHAEOBIOGEOGRAPHICAL ACCOUNT OF THE HISTORY OF THEIR
DISPERSAL ....................................................................................................................................................................... 112
5.1 INTRODUCTION ......................................................................................................................................................... 113
5.2 THE GRAIN FAUNA ................................................................................................................................................... 114
5.3 RECORDS OF THE FAUNA .......................................................................................................................................... 117
5.3.1 Pre-Roman Middle East ................................................................................................... 117
5.3.2 Prehistoric Europe ............................................................................................................ 119
5.3.3 Ancient Egypt .................................................................................................................... 120
5.3.4 Ancient Greece and Aegean ............................................................................................. 125
5.3.5 China ................................................................................................................................. 126
5.3.6 The Roman Period ............................................................................................................ 126
5.4 DISCUSSION: THE ORIGINS AND DIFFUSION OF THE SPECIES ................................................................................. 131
5.4.1 Neolithic ............................................................................................................................ 131
5.4.2 The Bronze Age ................................................................................................................. 136
5.4.3 The Iron Age ..................................................................................................................... 140
5.4.4 The Roman Empire ........................................................................................................... 142
5.5 CONCLUSION ............................................................................................................................................................. 147
CHAPTER 6 STABLE ISOTOPES (∆2H, ∆13C, AND ∆15N) FROM BEETLES ARE GEOGRAPHIC INDICATORS
OF THE ORIGINS OF CEREALS .................................................................................................................................. 149
6.1 INTRODUCTION ......................................................................................................................................................... 150
6.2 MATERIALS AND METHODS...................................................................................................................................... 150
6.2.1 Laboratory Rearing Experiment ...................................................................................... 150
6.2.2 Preparation of Chitin ........................................................................................................ 151
6.2.3 Removing Exchangeable Hydrogen ................................................................................. 151
6.2.4 Stable Isotope Analysis ..................................................................................................... 152
6.3 RESULTS .................................................................................................................................................................... 154
6.3.1 δ2H and δ13C Measurements from the Control Component ............................................ 154
6.3.2 Mixed Cereal Blind Tests δ2H .......................................................................................... 159
6.3.3 Buckwheat Blind Tests ..................................................................................................... 160
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6.4 DISCUSSION ............................................................................................................................................................... 160
6.5 SUMMARY.................................................................................................................................................................. 164
CHAPTER 7 THE APPLICATION OF ISOTOPIC ANALYSES TOWARDS INSECT REMAINS: MODERN
AND NEOLITHIC CASE STUDIES ............................................................................................................................... 165
7.1 INTRODUCTION ......................................................................................................................................................... 166
7.2 METHODS .................................................................................................................................................................. 166
7.3 CASE STUDY 1: WEST STOW, SUSSEX ................................................................................................................ 167
7.3.1 Site Information ................................................................................................................ 167
7.3.2 Collecting Methods ........................................................................................................... 167
7.3.3 The Fauna ......................................................................................................................... 167
7.3.4 Carbon Isotope Results ..................................................................................................... 168
7.3.5 Nitrogen Isotope Results ................................................................................................... 171
7.3.6 Hydrogen Isotope Results ................................................................................................. 172
7.3.7 Comparison of Isotopic Assays ......................................................................................... 173
7.3.8 Synopsis ............................................................................................................................. 176
7.4 CASE STUDY 2: NEOLITHIC GERMANY .................................................................................................................... 176
7.4.1 Site Information ................................................................................................................ 176
7.4.2 Collection Methods ........................................................................................................... 176
7.4.3 The Fauna ......................................................................................................................... 176
7.4.4 Carbon Isotope Results ..................................................................................................... 178
7.4.5 Nitrogen Isotope Results ................................................................................................... 181
7.4.6 Hydrogen Isotopic Results ................................................................................................ 182
7.4.7 Comparison of Isotopic Analyses and Discussion ........................................................... 183
7.4.8 Synopsis ............................................................................................................................. 188
7.5 CONCLUSION ............................................................................................................................................................. 189
CHAPTER 8 RECOVERY OF DNA FROM ARCHAEOLOGICAL INSECT REMAINS: FIRST RESULTS,
PROBLEMS AND POTENTIAL ..................................................................................................................................... 191
8.1 INTRODUCTION ......................................................................................................................................................... 192
8.2 MATERIALS AND METHODS...................................................................................................................................... 193
8.2.1 Thermal age: Accounting for Target Length and Copy Number .................................... 195
8.3 RESULTS .................................................................................................................................................................... 195
8.4 DISCUSSION ............................................................................................................................................................... 196
8.5 CONCLUSION ............................................................................................................................................................. 199
8.6 ACKNOWLEDGEMENTS ............................................................................................................................................. 199
CHAPTER 9 CONCLUSIONS AND FUTURE DIRECTIONS .................................................................................... 201
9.1 SUMMARY.................................................................................................................................................................. 202
9.1.1 The Palaeoecological Approach ....................................................................................... 202
9.1.2 The Biogeographical Approach........................................................................................ 204
9.1.3 The Isotopic Approach ...................................................................................................... 205
9.1.4 The Phylogeographic Approach ....................................................................................... 207
9.2 FUTURE DIRECTIONS ................................................................................................................................................ 208
9.3 CONCLUSION ............................................................................................................................................................. 209
BIBLIOGRAPHY .............................................................................................................................................................. 211
APPENDICES .................................................................................................................................................................... 250
APPENDIX 1UTUAL CLIMATIC RANGE DATA ................................................................................................................. 251
APPENDIX 2 EVALUATION OF BIOLOGICAL REMAINS FROM A ROMAN TIMBER DRAIN AT 21 ST PETERS STREET,
COLCHESTER (SITE CODE: 2007.124) ............................................................................................................................ 259
APPENDIX 3 STABLE-ISOTOPIC ASSAYS RELATING TO MODERN SITOPHILUS GRANARIUS, CEREALS, AND QUALITY
CONTROL PROCEDURES ................................................................................................................................................. 268
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APPENDIX 4 STABLE-ISOTOPIC ASSAYS FOR INSECT REMAINS RECOVERED FROM ERKELENZ-KÜCKHOVEN,
EYTHRA, PLAUßIG, AND WEST STOW ............................................................................................................................ 279
APPENDIX 5 RAW DATA FROM AMINO ACID RACEMIZATION (AAR) ANALYSES OF BEETLE REMAINS FROM
WATERLOGGED ARCHAEOLOGICAL CONTEXTS........................................................................................................... 283
APPENDIX 6 SUPPLEMENTARY DATA AND FIGURES PERTAINING TO THE GENETIC EXPERIMENTS DISCUSSED IN
CHAPTER 8 ...................................................................................................................................................................... 287
APPENDIX 7 TABLE 7.8...................................................................................................................................................................304
4
Acknowledgements
I would like to thank my supervisor Mr. Harry Kenward for the support he has given me
through each stage of my doctoral research.
His dedication and advice were invaluable to its
completion. He was also kind enough to share his work environment and welcomed the invasion of
beetle colonies into the laboratory. I am especially grateful for his unfailing encouragement as well as
his unflappable willingness to always go the extra mile in support my endeavours.
I would also like to extend my appreciation to Dr. Allan Hall and Dr. Matthew Collins who
offered valuable suggestions to improve my research. Moreover, Dr. Hall supplied a multitude of
environmental samples for processing, and he was able to endure mishaps in the lab with a smile. Dr.
Collins served as a catalyst for experiments conducted in association with the University of
Copenhagen, the University of Glasgow, and Bradford University.
Dr. Tom Gilbert deserves a special mention, for without his patience and help I would never
have been able to understand the procedures necessary to explore the potential of the genetic
methodology. All the members of the laboratory in the Department of Ancient DNA and Evolution at
University of Copenhagen were willing to help and address questions. My thanks extend to all of
them.
A number of people and organisations assisted in providing specimens for this project. In
particular, I would like to acknowledge Edith Schmidt for donating archaeological specimens from the
German sites and Philippe Ponel for providing archaeological and modern individuals from sites in
France. Laible Friedman of the Department of Zoology, Tel Aviv University and the team from
Central Science Laboratories deserve my thanks for providing numerous grain beetles from their
respective museum collections. In addition, I am indebted to Laura Girven and Melanie Rousseau for
their assistance processing and sorting the archaeoentomological samples.
I wish acknowledge the Palaeoecology, Archaeology, and Evolutionary Origins (PALAEO)
research group for granting the Marie Curie Early Stage Short-term Training Fellowship. I am also
grateful to the Department of Archeology, University of York for bestowing the Innovation and
Research Priming Grant. The funding assisted with equipment fees, experiment costs, and travel.
Special thanks go to Emma Bradley and my parents, Gary and Bonnie King, for enduring
cheerfully through the emotional roller-coaster of living with someone involved in such as long-term
endeavour. They kept my spirits up through offering encouragement and advice and sometimes just
by lending an ear.
5
Chapter 1
Introduction and Scope of Work
6
1.1. Trade and Migration: The Archaeological Viability
Throughout time, humans have migrated and interacted, and thus have both purposely and
unwittingly carried animal species with them in their travels. While a few of the human-introduced
animal species were able to permanently colonise these new geographical areas, numerous species
failed to find suitable niches due to environmental or climatic restraints. The ability to distinguish the
native species from the immigrants is archaeologically significant. However, the mapping of animal
movement is not an end in itself rather it provides insight into the movement and interaction of past
peoples. Additionally, the presence of man and the introduction of new animal species into a preexisting ecosystem may lead to local extirpations, such as the pygmy hippopotamus in Cyprus
(Simmons 1988) or the giant lemurs of Madagascar (Perez et al. 2005), and likewise the desertion of a
region by man may have detrimental effects on strong synanthropes (Brothwell and Jones 1978).
Given this, the study of faunal remains may shed light on past exchange and trade networks (e.g.
Hodges 1982; Iregren 1988) as well as migration and population movement (Ashby 2004).
The ability to discern past exchange and trade patterns is archaeologically valuable. Exchange
is an inclusive concept, which may be viewed as the spatial distribution of material items, ideas, and
information between individuals and social groups (Earle 1982) with the payment being either
immediate or delayed and indirect (Alden 1982). However, trade is a more narrowly defined and
archaeologically visible index of exchange (Crabtree 1990) that refers specifically to material goods
(Roslund 1992). The concepts are archaeologically interesting because commentators have postulated
that exchange and trade were impetuses for urban and social development and cultural change
(Hodges 1982; Wells 1980). With knowledge of exchange and trade routes, inferences may be made
concerning economic conditions, population pressures and settlement patterns (Hodges 1982).
Human movement through migration is another deep concern of archaeology. Although the
archaeological visibility and legitimacy of migration have been questioned in the past (Clark 1966;
Härke 1998), it has recently experienced resurgence (e.g. Anthony 1990, Burmeister 2000, Barrett et
al. 2001). There are two main types of migration: immigration, a small-scale event, and population
movement, a large-scale phenomenon.
Migration as a small-scale event is difficult to infer
archaeologically because the immigrants are assimilated into the local population (Burmeister 2000).
However, population movement is made evident by the ensuing cultural change resulting from the
absorption or forcing out of the local population by a large and powerful migrating body (Rouse
1986).
In past studies, the traditional methods for the identification of culture contact have too often
resulted in the assumption that trade was the primary or sole mechanism for cross-culture interactions
(Olausson 1988). The traditional methods include: polysemous contextual analysis (e.g. Hodder
1982), stylistic analysis (e.g. Lindstrom and Kristoffersen 2001), identification of raw materials (e.g.
7
Rosenfeld 1965; Galloway et al. 1996), spatial distribution, and the presence or absence of a local
precedent (Olausson 1988).
Ethnographic comparisons have demonstrated that when applied
individually, the traditional methods have severe limitations derived from assumptions that the
imported objects will be:
•
foreign in origin;
•
valuable;
•
unavailable in the local environment;
•
limited in number (Olausson 1988).
These problems largely stem from the framework for analysis of culture contact and ethnicity
being based in culture history, which unfortunately results in the identification techniques for foreign
objects relying heavily on overly simplistic regional sequences of artefact development (Jones 1997).
As Wells (1980) mentioned, style may signal acculturation, and the differences in function risk being
mistaken for ethnic variation (Jones 1997). As archaeological cultures are not absolutes, direct
correlations cannot be drawn with ethnic units (Jones 1997). It may be more effective to approach the
study of culture contact through an index that reflects social and cultural relations but is less dependent
on arbitrary cultural distinctions, i.e. ethnic expression (Jones 1997).
One method of countering these innate problems is through utilisation of holistic approaches.
For example, stylist comparisons should be applied in addition to identification of raw materials
(Hantman and Plog 1982) and contextual analyses (Hodder 1982). However, there are still other
distortions in the interpretation of culture contact—it is disproportionately based on grave goods (e.g.
Wells 1980) and may be skewed by artefact recovery based on the types of sites. This is made evident
by studies of the Roman exchange in Europe, which are biased towards maritime trade because of the
dominance of recovered amphora in the contexts (Pentz 1992). Faunal analysis is a promising area,
which if applied to the study of human movement, would balance the evidence and avoid arbitrary
cultural distinctions (Ashby 2001).
Ashby (2004; 2006) has approached the question of exchange and population movement from
a zooarchaeological standpoint. In archaeology, animals are typically viewed in association with
production and consumption rather than the realm of exchange (Earle 1982). However, details of
culture contact may be extrapolated using standard zooarchaeological methods:
•
metric (continuous) variation;
•
non-metric (discontinuous) variation;
•
genetic analysis;
•
species biogeography (Ashby 2004).
Metric and non-metric traits are beneficial for recognition of genotypes, age and sex of
individuals, and bone shape variation (O’Connor 2000). Through examination of these variations, it is
8
possible to ascertain information about phylogeny (e.g. Shigehara et al. 1993), the human factor in
selection (e.g. O’Connor 2001), and trade (e.g. Murphy et al. 2000). Genetic work on the Pacific rat,
Rattus exulans, has demonstrated the potential of DNA analysis in determining relationships between
populations and thus provides insight into culture contact (Matisoo-Smith and Allen 1997; 2001;
Matisoo-Smith and Robins 2004).
Biogeography is essential to the understanding of animal
movement (e.g. Barrett 1997) because it aids in the recognition of foreign species and may shed light
on their original geographic region.
However, a zooarchaeological approach to culture contact is not without problems, notably
taphonomy and inter-analyst variability. It is essential to consider the transposition of the living
community into the archaeological assemblage, which is known as the taphonomic process (Efremov
1940). The final sampled and interpreted assemblage consists of the preserved remains of both
autochthonous and allochthonous components. As the biotic assemblage proceeds through the various
taphonomic stages, the ecological signal becomes distorted from its original state as increasingly more
information is lost. The majority of taphonomic effects are beyond the control of the researcher, and
the best that one may do is account for them in the reports. However, problems with artefact recovery
and inter-analyst variability are manageable through the standardization of methods and exercise of
caution.
Although not entirely void of problems, the archaeological study of faunal remains and animal
movement has proven capable of allowing inferences into identity and culture contact (e.g. Ashby
2006). An excellent example of the benefits of zooarchaeological analysis is illustrated by Rausling’s
examination of the presence of Camelus bactrianus bones in 11th century BC Mesopotamia (Rausling
1988), which implies a westward connection 1000 years before it is evidenced in documentary
accounts.
While the applicability of animal remains to studies of culture contact has been investigated,
most of the assessments have been restricted to vertebrate zooarchaeology with little attention given to
invertebrate species. This is surprising given the strong ecological signal demonstrated by some
insects (making them prime candidates for biogeographical study), the direct evidence that foreign
product-associated species were transported on ships (e.g. Pals and Hakbijl 1992), and the discovery
of import indicator species from archaeological sites (e.g. Osborne 1971). This manuscript will focus
on employing insect fossils for the purpose of inferring human movement and culture contact.
1.2. The Archaeological Significance of Insects
While insect fossils have been noted in archaeological sites for over a century (e.g. Roeder
1899; Bayford 1903), most of the advancements in the field of archaeoentomology have occurred
since the 1970s. Insects have proven to be immensely valuable for the reconstruction of past climates
9
(e.g. Coope et al. 1998) and environments (e.g. Hill 1994b). They have been employed in spatial
reconstructions of structures (e.g. Buckland et al. 1983) and interpretations of living conditions (e.g.
Panagiotakopulu 2001). Recently, there has been a developing interest in the use of insects as
archaeological indicators of human activity.
Reconstructing the level of human impact on an environment may be problematic especially
after the onset of the Roman period (see Kenward in press). This is largely the result of the paucity of
insect fossils collected from natural settings (e.g. Hill 1994b); however, sampling methods and
taphonomy are also factors. Despite these caveats, Carrott and associates (1995a) have investigated a
‘natural’ layer beneath a circa 12th century strata at Keldergate, Beverly, which implies an
occupational expansion into the area during that period. Also in the Period 3 samples (Hall and
Kenward 1999a) from 16-22 Coppergate, York, there is evidence to suggest that the area was
unoccupied (though used for rubbish deposits) prior to the construction of Anglo-Scandinavian
tenements in Period 4 (Hall and Kenward 1999b; Kenward and Hall 1995). Some of the best evidence
of human influence on the environmental is derived from the remains of drains, moats, and artificially
constructed ponds. The invertebrate remains from the Higher Lane, Fazakerley, Merseyside site
(Dobney et al. 1995) demonstrate the transition from the initial construction of an aquatic arena, to a
stable aquatic body, and finally to a terrestrial area.
The archaeoentomological field of human
environmental impact is still in its infancy yet if properly applied, provides insight into human activity
that may not be visible through traditional, material culture based approaches.
Additionally, archaeoentomology has demonstrated great potential in unveiling the human
exploitation of resources through providing secondary evidence for the presence of livestock, cereals,
vegetables, and other raw materials. Through scrutiny of the ecology of the various insect species, it is
possible to ascertain insect associations with certain materials that would have been utilised by the
humans.
At 16-22 Coppergate, York, the insect remains provide evidence for the availability of a
number of resources. For example, the bark beetle Leperisinus varius was identified in forty-eight
contexts (Kenward and Hall 1995) and indicates, through its ecological association, the presence of
Fraxinus (ash) at the site. Indeed, this claim is supported by the recovery and identification of ash
wood in 275 records from 16-22 Coppergate (Kenward and Hall 1995). At Nipáitsoq, Greenland
(Buckland et al. 1983), the presence of taxa like Byrrhus fasciatus and Simplocaria tesselata imply
that moss was employed at the site, possibly for floor-layers, bedding, or sanitary paper. When used
in conjunction with other palaeoecological methods, the invertebrate remains provide another layer of
evidence to substantiate the presence of certain resources.
Hall and Kenward (2003) have reviewed the potential of environmental remains as indicators
of crafts and industries such as tanning, wool-processing, and dyeing. It has been postulated that
insect remains may be used as signs of the tanning industry. Trox scaber, Acritus nigricornis,
Creophilus maxillosus, Teretrius fabricii, and Phymatodes testaceus have all been suggested as a
10
probable indicator group for tanning (Kenward in press). However, this association is tentative, and
Hall and Kenward (2003) caution about using the species alone to discern tanning. Another craft
indicator is the sheep ked Melophagus ovinus, which lives in wool, and has been argued as evidence
for areas of wool production (Buckland and Perry 1989). Furthermore, the plant dyer’s greenweed
(Genista tinctoria) is believed to have been exploited in Anglo-Scandinavian York for its use as a dye.
The existence of the dye industry is further supported at the 16-22 Coppergate site by the presence of
the weevil Apion (Exapion) difficile, which is strongly associated with dyer’s greenweed (Kenward
and Hall 1995). The discovery of Apion difficile is more interesting still because it is rare amongst the
list of British beetles and is likely to be representative of another human activity, trade.
1.3.
Aims of the Study
Studies of biological evidence from archaeological sites are providing a wealth of information
about palaeoecology and human activities. Many commentators have noted that faunal remains are
important and useful elements for inferences concerning resource exploitation, production,
consumption, and industries. Recently, Ashby (2006) has demonstrated that an examination of faunal
remains such as bone and antler may be used to ascertain information about identity and culture
contact in the Viking Age. While the prospect of insect fossils as valuable indicators of culture
contact has been proposed (e.g. Sadler 1988; Kenward in press), its potential has yet to be sufficiently
addressed.
The few attempts to connect insect transportation to human movement have been
constructed on biogeographical grounds within the documented historic period (e.g. Lindroth 1957;
Hammond 1974). However, these studies fail to provide a systematic index for the identification of
transported insects or a means to substantiate their point of origin. In the present text, a systematic
exploration of potential archaeoentomological approaches for discerning culture contact is undertaken.
1.4.
Approaching a Methodology
Culture contact cannot be fully understood until its medium has been established. Thus, the
first phase in analysis is to develop an index for determining which archaeoentomologically
identifiable products were being distributed. While humans may have been engaging in the exchange
of insect products (e.g. honey; silk), knowledge of the ecological requirements of insect species
provides secondary evidence of the human exploitation of resources [Section 1.2]. Therefore, the
medium for culture contact can be established directly through evidence of the insect products
themselves and indirectly through product-associated insect species.
Once the exchangeable products have been identified, it is necessary to determine their source.
Insects may offer evidence of both short (at a local or regional level) and long-distance trade.
Knowledge of the source of the resources is essential to developing an understanding of culture
11
contact. Are the materials foreign in origin or locally accessible? If they were locally available, were
the inhabitants electing to import the resources long distances? Why were the products being
exchanged?
An understanding of insect morphology, physiology, and ecology is beneficial in
distinguishing some of the foreign (alien) species from the natives or colonists.
Through
reconstruction of past climates and environments, the palaeoecology for a site can be established
[Chapter 4]. Thus, it is possible to determine whether a species would be capable of surviving in a
past ecosystem—ecological outliers being representative of imports. While this method serves to flag
a few of the imported species, Lindroth (1957) and later Hammond (1974) have shown the problems
of this method when sorting the natives from the aliens, and alternative approaches need be
considered.
There are three main tools available to archaeoentomology which can be employed to trace the
movement of faunal materials: species biogeography, isotopic analysis, and phylogeography. Species
biogeography [Chapter 5] is essential to understanding animal movement without relying on chemical
or genetic techniques (Barrett 1997; Buckland and Sadler 1989). Biogeography notes diachronic
changes in species distribution as well as geographical ranges and thus allows for the tracking of
animal movement. Although stable isotope analysis [Chapter 6; Chapter 7] has largely been employed
in climate studies (Schimmelmann and DeNiro 1986; Gröcke et al. 2006), it may be used to identify
autochthonous and allochthonous components within and between sites. The oxygen isotope ratio,
18
O/16O, in meteoric water is quite regionally distinctive and may become fixed in the tooth enamel or
chitin-layers of the inhabitants of those regions. Oxygen isotope analysis thus reveals the regional
signal of the place in which the individual was living at the time of the chitin or enamel layer was
formed. Hydrogen isotopes, D/H, function in the same manner as oxygen isotopes; however, they
reflect hydrogen that was ingested as organic hydrogen or water hydrogen (Gröcke et al. 2006). By
examining the isotopic signals preserved in alien chitin, it may be possible discern the origin of the
associated imports.
Phylogeography [Chapter 8] is the study of genetic relationships among
populations of species, which sheds light on the length of time certain populations have been isolated
from each other (e.g. Smith and Farrell 2005; Moya et al. 2004), and as such, serves as an index for
discerning past culture contact.
It is the aim of this study to define a methodology which is archaeologically applicable
regardless of temporal or geographic boundaries. However, a comprehensive study of insect remains
from all archaeological contexts is not feasible. Instead, case studies will be selected from the United
Kingdom and Continental Europe.
12
1.5.
Structure
While culture contact has been investigated utilising material culture and zooarchaeology, it
has yet to be studied in an archaeoentomological framework. This book is divided into nine chapters.
Following this brief introduction, Chapter 2 will examine beetle morphology and ecology.
A
discussion of these issues is essential as they form the basis for any exploration into
archaeoentomology. Form and function are the prerequisites for the success of an organism in its
environment (Speight et al. 1999) and are essential to understanding an insect’s ecological role. As
such, Chapter 2 will discuss the concepts of habitats, ecosystems, and ecological associations. Elton
(1927; 1966) examined several types of interactions between the same species, species of the same
ecological niche, and species of different trophic levels.
Through scrutiny of these ecological
associations, it will be possible to ascertain connections between certain insect species and humanexploited materials or ‘products’. Furthermore, palaeoecological factors like climate and environment
will be employed to pin-point alien, foreign, species, which would have been unable to survive in the
local area due to ecological constraints, and separate them from the native or successfully colonial
species.
Having established the biological and ecological framework [Chapter 2], Chapter 3 will outline
the methodological approaches taken in the project—palaeoecology, biogeography, isotopic analyses,
and phylogeography. In addition to the information provided in Chapters 2 and 3, these methods are
illustrated primarily through the use of case studies.
The techniques are applied in Chapters 4-8. Chapter 4 considers the insect remains from two
case studies—Roman 7-15 Spurriergate, York and Anglo-Scandinavian 16-22 Coppergate, York—
through a palaeoecological approach. The palaeoecological approach is employed to demonstrate the
ability of archaeoentomological remains to stand as secondary evidence of commodities and materials
that were used by humans in the past. Chapter 4 uses palaeoecology as a tool towards inferring culture
contact. Chapter 5 presents a biogeographical investigation of human migration and culture contact as
evidenced through grain-associated insect species. In Chapters 6 and 7, stable-isotopic analyses of
carbon-13, nitrogen-15, and deuterium are reviewed from modern and Neolithic insect specimens.
Chapter 8 provides evidence towards the applicability of genetic analysis to insect fossils and
discusses the potential of phylogeography as a palaeoeconomic tool. Following the presentation of the
case studies and the results of the methodologies, their significance and meaning are discussed. While
the functionality and practicality of the various methodologies are assessed, the emphasis is on culture
contact. The text closes with general conclusions and comments concerning the applicability of
archaeoentomology to culture contact and the potential for future research.
13
Chapter 2
Morphological and Ecological Concepts
14
2.1 Introduction
The formulation of hypotheses constructed from insect fossils is dependent upon the validity of
the existing knowledge of the species
species.. In this chapter, the published methods for insect species
identification will be reviewed; followed by a discussion of the ecological princ
principles
iples inherent in the
archaeological and palaeoenvironmental interpretations of the fossils.
This presentation of the
morphological and ecological concepts will form the basis for any archaeoentomological argume
arguments
that arise during the study as well as eestablishing the biological background for exploration of
biogeography, isotopic analyses, and phylogeography.
2.2 Morphology
While this work is not primarily concerned with the form and structure of insects, a brief
review of insect morphology is worthy of consideration because:
• it plays a significant role in the identification methods employed by both modern entomologists and
palaeoentomologists;
• the ability of an organism to succeed in its environment is dictated by form and function
function; and,
• the biosynthesis involved in the formation of specific morphological components is a function of its
environment.
Insects are six-legged,
legged, segmented inv
invertebrates that possesss the arthropod’s characteristic
articulated, external skeleton (i.e. exoskeleton). Taxonomic recognition of insect orders, families, and
genera is often established through examination of the myriad anatomical features of the
appendages—mouthparts,
mouthparts, legs, wings, and abdominal apex, and moreover, species are almost
exclusively denoted based on anatomical distinctions. Because of the multitude of variations, it is
only possible here to provide a basic overview of the external anatomy.
The basic structure of a typical adult insect [Figure 2.1] consists of three
Figure 2.1 Insect Morphology
(Borror and White 1970, 30)
specialised sections called tagmata—a 6-
Figure 2.2 Types of Body Segmentation
segmented head for sensory perception and
food gathering, a 3-segmented thorax for
locomotion, and an 11-segemented abdomen
for digestion and reproduction (Speight et al.
1999). While these external features have been
amalgamated into functional units in adult and
nymphal insects, the segmentation evidenced
in the sclerotized adults and nymphs is not
directly
homologous
unsclerotized
larvae
with
[Figure
that
2.2],
of
the
which
possess the distinctive metameric segmentation
apparent in annelids (Gullan and Cranston
2000).
In
adults
and
nymphs,
the
sclerotization stretches beyond the primary
segment, commencing in front of the fold and
A) Primary segmentation evinced by soft-bodied
larvae; B) Simple secondary segmentation; C) More
derived secondary segmentation (Snodgrass 1935)
extending almost to the rear of the segment,
which allows the muscles in the folds to attach to solid rather than soft cuticle (Gullan and Cranston
2000). In defined areas, the sclerotization process produces plates known as sclerites—the tergum
(dorsal plate), the sternum (ventral plate), and the pleuron (side plate). In the head, all the sclerites are
fused together into a rigid capsule. However, the abdominal pleura remain partially membranous, and
on the thorax, they are sclerotized and linked to the tergum and sternum of each segment. During the
sclerotization process, the progressive hardening of the proteinaceous structures is utilised by insects
to stiffen and harden their exocuticle (Hopkins and Kramer 1992).
2.2.1 The Cuticle
The cuticle of insects functions both as an exoskeleton and a barrier between the living tissues
and the environment and, to a large extent, determines the shape and appearance of insects. This thin
layer of material consists of two main strata, a thin (0.1- 3 µm), outer epicuticle and a thicker (0.5- 10
µm) inner procuticle, which are secreted from an underlying single layer of cells, known as the
epidermis [Figure 2.3]. As the outermost layer, the epicuticle serves to prevent dehydration and block
invasive foreign matter. It consists of three layers, of which the innermost stratum, the cuticulin layer,
16
is composed of lipoproteins and chains of fatty acids embedded in a protein-polyphenol complex. A
monolayer of wax molecules, which are bipolar (i.e. with hydrophilic and hydrophobic ends), resides
above the cuticulin layer and serves as the chief barrier to water passage in and out of the insect’s
body. In many insects, a third cement layer protects the waxy layer from abrasion and heat loss.
Whereas the epicuticle acts as a barrier against external factors, the procuticle, which consists of a
thick, pale endocuticle and a thin, often dark exocuticle, provides structural support. The procuticle is
made primarily of chitin chains; its two layers being differentiated by the sclerotization of the
exocuticle (Gullan and Cranston 2000).
While thin and flexible in many larvae, the adult cuticle is typically both rigid and armour-like
(e.g. mandibles) and tough and elastic (e.g. abdomen) depending on its function and location in the
body. The strength of the cuticle is derived from the extensive hydrogen bonding of adjacent chitin
chains. Chitin is an amino-sugar polysaccharide predominantly composed of β (1-4) linked units of
N-acetyl-D-glucosamine (Gullan and Cranston 2000). Further rigidity is established in the exocuticle
through the irreversible process known as sclerotization, which contributes to the proteins becoming
water-insoluble. In the membranes between joints or segments, a solid cuticle is not conducive for the
necessary flexibility, and an unsclerotized soft cuticle containing resilin is formed instead with a
thicker endocuticle and a thinner, or absent, exocuticle. While the rigidity of the exocuticle is
essential in certain parts of the insect’s body, the benefit of a water-soluble soft cuticle is evidenced by
the abdominal dilation permissible in the worker honeypot ant, Camponotus inflatus, which retains
honey in its distensible abdomen as a colonial food store (Hadley 1986).
The procuticle lies directly on the epidermis, which is the single-celled stratum underlying the
cuticle that is responsible for the production of the cuticular components, waxes, cements, and defence
mechanisms (Gullan and Cranston 2000). The epidermis secretes cuticle through an organised array
of microvilli upon its apical face. At the top of the epidermal microvilli, the cuticulin is deposed
which then extends to form a continuous envelope. Through a different cellular process, the epicuticle
is formed from the secretory discharge of epidermal vesicles (Payre 2004). Locke (2001) postulates
that the laminae of chitin microfibers composing the procuticle are developed in an assembly zone
located above the microvilli.
The epidermal secretion process is the impetus in both the
transformation and preservation of once living cellular material into the tough, durable cuticular
layers.
It is the robustness of the cuticle layers that directly correlates to an insect’s preservability in
archaeological contexts. Therefore, Coleoptera (beetles) specimens are frequently archaeologically
recovered due to the pronounced hardiness of their cuticles whereas Lepidoptera (butterflies and
moths) are rarely present. Sadler (1988, 14) claims that during the formation of the thanatocoenosis
stage, “the soft parts of all insect families are lost… [and] in the poorly sclerotized groups, however,
such as, the Lepidoptera, Diptera, Hymenoptera and many of the Anoplura and Siphonaptera, the
17
majority of the animal is lost”. Additionally, it is the more sclerotized portions, i.e. the head, the
thorax, and the elytra, of the coleopteran specimens that survive taphonomically.
While the
preservation of morphological characteristics is crucial in the identification process, the survivability
of Coleoptera, which do not moult in their adult stage, suggests that a cellular record of their complete
adult life is retained in their subfossil remains allowing for the recovery of and experimentation with
viable amino acids, isotopes, and ancient DNA.
Figure 2.3 The Cuticle
(Hadley 1986)
2.2.2 The Chitin
Chitin is the principle structural component of the insect body and accounts for approximately
25-40 % of the dry weight of insect cuticle (Jeuniaux 1971). It consists predominantly of unbranched
18
chains of β-(1, 4)-2-acetamido-2-deoxy-D-glucose (i.e. N-acetyl-D-glucosamine) and occurs in the
procuticle but is absent from the epicuticle. Hackman (1964) suggests that chitin may be viewed as a
derivative of cellulose in which the hydroxyl groups of the second carbon of each glucose unit have
been replaced by acetamido (-NH (C=O) CH3) groups.
During the pharate phase (the period of moulting when an insect has secreted a new cuticle but
has not escaped the old cuticle) and immediately after ecdysis (the shedding of the integument) has
taken place, chitin is synthesized most rapidly (Chippendale 1978). The chitin is produced in the
epidermal cells from the sugar nucleotide, UDP-2-acetmido-2-deoxy-D-glucose.
According to
Chippendale (1978), the synthesis of chitin from glucose in insects involves the series of reactions
presented in Table 2.1.
Table 2.1 Steps leading to chitin synthesis
1.) ATP + D-glucose
=
2.) D-glucose 6-phosphate
=
3.) D-fructose 6-phosphate +
L-glutamine
=
4.) Acetyl-CoA + 2-amino-2- deoxy- =
D-glucose 6-phosphate
5.) UTP +2-acetamido-2-deoxy-D- - =
Glucose 1-phosphate
6.) UDP-2-acetamido-2-deoxy-D=
glucose + [1, 4-(2-acetamido-2-deoxyβ-D-glucosul)]
ADP + D-glucose 6-phosphate
(hexokinase)
D-fructose 6-phosphate
(glucosephosphate isomerise)
2-amino-2-deoxy-D-glucose 6-phosphate
+ L-glutamate (glucosaminephosphate
isomerise)
CoA + 2-acetamido-2-deoxy-D-glucose
6-phosphate (glucosaminephosphate
acetyltransferase
pyrophosphate + UDP-2-acetamido2-deoxy-D-glucose (UDPacetylglucosamine pyrophosphorylase)
UDP + [1, 4-(2-acetamido-2-deoxy-β-Dglucosyl)] (chitin synthase).
The metabolic reactions involved in the biogenesis of chitin directly impact the isotopic
analyses that contribute to this research endeavour. The photosynthetic production of glucose, the
building blocks of carbohydrates, in plants is a combination of atmospheric CO2 and environmental
water (i.e. precipitation).
6 CO2 + 6 H2 O C6H12O6 + 6 O2 [Formula 2.1]
Insects feeding on the plants, or plant products, ingest dietary carbohydrates, such as starch,
which in turn are hydrolyzed by the enzyme endoamylase (1, 4-α-D-glucan glucanohydrolase). The
enzyme attacks the interior glucosidic bonds of the starch producing a mixture of linear and branched
oligosaccharides. The hydrolysis of the starch to absorbable glucose is completed in the intestine
using the enzymes endoamylase, α-glucosidase, and oligo-1, 6-glucosidase. Afterwards, the nutrients
are absorbed from the lumen of the intestine to the hemolymph through a rate that is controlled by two
known limiting factors:
19
1.) the rate of release of fluid from the crop into the midgut;
2.) the rate of conversion of absorbed glucose into trehalose, i.e. a storage and transport sugar
(Chippendale 1978).
In the insect’s body, the carbohydrates are stored as glucose, glycogen, or trehalose until used
[Figure 2.4]. In regards to the chitin synthesis, the glucose is most likely the derivative of a trehalose
precursor. During the biogenesis of chitin, hydrogen can be added to the carbon ring during two steps
[see Table 2.1].
In step 3, the amino acid L-glutamine combines with D-fructose 6-phosphate
resulting in 2-amino-2-deoxy-D-glucose 6-phosphate and permitting the NH3 amino group to bond
with the second carbon ring in order to add two new hydrogen atoms. In step 4, acetyl CoA (note that
the hydrogen of acetyl CoA originates from glucose-derived pyruvate via the Embden Meyerhoff
pathway) reacts with the amino group:
NH3 + O
S –CoA NH + S –CoA
C
C=O
CH3
CH3
[Formula 2.2]
Of the ten carbon-bound hydrogen atoms contained in one unit of chitin (C8H13NO5), seven
were derived from the glucose produced during photosynthesis and three from pyruvate. The
nitrogen-bound hydrogen is obtained from the L-glutamine, leaving two hydrogen atoms which are
derived from an unaccounted source (Miller 1984).
Figure 2.4 The interactions of glucose, trehalose, and glycogen
(A) hexokinase (inhibited by glucose 6-phosphate; (B) glucose-6-phosephatase (activated by trehalose); (C)
phosphoglucomutase; (D) glucose-1-phosphate uridylytransferase; (E) glycogen synthase (activated by
glucose 6-phosphate; (F) glycogen phosphorylase (inhibited by ATP); (G) α, α-trehalose-phosphate
synthase (activated by Mg 2+ , inhibited by trehalose); (H) trehalose phosphatase; (I) α, α-trehalase; (J)
glycolytic enzymes (control at phosphofructokinase and pyruvate kinase)
(Redrawn from Sacktor 1970 in Chippendale 1978, 33)
20
2.2.3 The Head
The head is the rigid capsule-like structure that constitutes an insect’s anterior body region.
The cranial capsule has only two openings, one to the mouthparts and the other through the occipital
foramen to the prothorax. It is composed of six fused segments:
• labral;
• antennal;
• postantennal;
• mandibular;
• maxillary;
• labial (Gullan and Cranston 2000).
An insect’s head is host to its mouthparts, eyes, and antennae. The mouthparts are constructed
from all the head segments except the antennal, consisting of the upper (labrum) and lower (labium)
lips, the jaws (mandibles), and two jaw-like appendages (maxillae) (Knopf 1980). Additionally,
Gullan and Cranston (2000) list the hypopharynx, which is a tongue-like feature, amongst the basic
mouth components. While the labrum and the labium form a preoral cavity, the hypopharynx divides
the cavity into a food pouch and a salivarium. The mandibles and the maxillae assist in the processing
of food. While the mandibles, which may possess an indentation hardiness of up to 30 kg mm-2 and a
3 on the Moh mineral hardness scale, cut and crush food, the maxillae hold and macerate the food
(Gullan and Cranston 2000).
In addition to functioning as food acquisition and processing tools, insect mandibles feature
prominently both as defence mechanisms and instruments for sexual selection. In some beetle species,
like Lucanus cervus, the mandibles are analogous to antlers in elk and are similarly larger in males and
employed by them in fights over females (Preston-Mafham 2005). Evolutionary adaptations for food
processing, defence, or sexual selection have produced myriad mouthpart types based upon the basic
components and design.
These modifications to mouthpart structures are exploited by phenotypic identification
methods and serve as the foundation for ecological generalisations. The classification of mouthparts is
typically categorised by feeding method [Figure 2.5] chewing, sucking, or piercing (Borror and White
1970). Chewing mouthparts are typically characterised by laterally moving mandibles that cut and
crush the food; however, certain insects such as bees evince a chewing and lapping method. Gullan
and Cranston (2000, 27) define lapping as “a mode of feeding in which liquid or semiliquid food
adhering to a protrusible organ, or ‘tongue’, is transferred from substrate to mouth.” In bees, the
mandibles are restricted to manipulation of wax, fighting, feeding larvae, and labour (Gullan and
21
Figure 2.5 Types of Mouthparts
(Knopf 1980, 16-17)
22
Cranston 2000). Most adult Lepidoptera (moths and butterflies) have evolved suctorial mouthparts in
the form of an elastic proboscis, which pumps liquid food. While various modifications are indicative
of piercing mouthparts, they largely consist of needle-like stylets or proboscis which specialise in
breaking animal and plant tissue (Borror and White 1970). Afterwards, the piercing insects acquire
their food through sponging the pre-existing liquids, e.g. Anoplura (sucking lice), or by extra-orally
digesting any solids and sucking the resulting liquids, e.g. larval Neuroptera (net-winged insects), into
the food canal (Gullan and Cranston 2000).
Figure 1.6 Types of Antennae
In addition to mouthparts, most insects have
two kinds of eyes—simple and compound. Simple
eyes, termed ocelli, are merely sensitive to light and
are not designed for high-resolution vision (Knopf
1980). Generally, the ocelli reside in the triangular
portion on the top of an insect’s head, permitting the
insect to respond to subtle changes in light (Gullan
and Cranston 2000).
The compound eye is the
insect’s most sophisticated visual organ, allowing
coverage of nearly 360 degrees of space. Compound
eyes are comprised of several minute lenses which
provide the insect with panoramic images formed
from apposed points of light (Gullan and Cranston
2000). Further sensory structures exist in the form of
paired
mobile,
segmented
appendages
called
antennae, which insects use for smell, touch, and
occasionally hearing (Knopf 1980). The antennae
contain a multitude of hairs, pits, and cones that
operate
as
sensory
organs
in
the
form
of
chemoreceptors, mechanoreceptors, thermoreceptors,
and hygroreceptors (Gullan and Cranston 2000).
Size, shape, and elaborateness vary between insects
[Figure 2.6] and can be indicative of taxonomic
groups. Moreover, antennae may often be examined
(Knopf 1980, 14)
to ascertain gender because male insects may have more elaborate antennae than females (Gullan and
Cranston 2000).
The morphological variations of insect head features, especially the mouthparts and antennae, are
often exploited for entomological identifications.
As one of the more commonly recovered
archaeoento-mological fragments, the distinct-iveness of the fossilised heads is often beneficial in the
determination of taxonomic order and genera. However, analysis of thoracic and/or abdominal remains
is occasionally necessary for species identification.
2.2.4 The Thorax
The thorax is made up of three segments: the prothorax, the mesothorax, and the metathorax.
Each thoracic segment has one pair of six-segmented legs; in the case of winged insects, the
mesothorax and usually the metathorax bear wings (Knopf 1980). The thoracic tergal plates are
variously modified in pterygotic adults though remain rather simple in apterygotes and immature
insects (Gullan and Cranston 2000). In beetles, mantids, grasshoppers, and some Orthoptera, the
upper surface of the prothorax, the pronotum, is large, extending from the head to the wing base, and
in cockroaches, it serves as a shield that covers part of the head and mesothorax (Knopf 1980).
Because of its uniqueness, the preservation of the pronotum in the fossil record assists in the
palaeoentomological identifications.
Insects have three pairs of legs—fore, mid, and hind—composed of five parts: the coxa, the
trochanter, the femur, the tibia, and the tarsus (Knopf 1980). The apex of the tarsus usually wields a
pair of claws and pad-like structures (Borror and White 1970). While the femur and the tibia are
generally the longest leg segments, variations exist based on function (Gullan and Cranston 2000).
These functional modifications may be indicative of behavioural specialisations such as the enlarged
forelegs of mantids for snaring prey and the developed hind femora and tibiae of grasshoppers
designed for leaping (Knopf 1980).
Further specialist adaptations are made evident by the thoracic wings. Although all winged
insects are believed to share a homologous eight veined venation ground plan, venation patterns are
only consistent within groups and further wing variations exist in shape, size, and degree of
sclerotization [Figure 2.7] (Gullan and Cranston 2000). In the case of Coleoptera (beetles), the
forewings have hardened into protective wing cases, elytra, which secure the hind-wings when an
individual is not in flight [Figure 2.7d]. Another wing modification is apparent in the Diptera (flies)
[Figure 2.7 f], in which the hindwings have been adapted as a balancer (Gullan and Cranston 2000).
In adult insects, studying the positioning of wings helps distinguish between groups. For example,
some members of Heteroptera and Coleoptera encase their membranous hindwings in their thickened
forewings (i.e. elytra) when they are held flat against the back (Preston-Mafham 2005), suggesting
convergent evolution between the taxonomic orders. However, in Heteroptera, the membranous wing-
24
Figure 2.7 Wing Modifications
(a) Forewing of a butterfly; (b) Forewing of a dragonfly; (c) Forewing of a cockroach; (d) Forewing of a beetle;
(e) Forewing of a mired bug; (f) Forewing and haltere of a fly
(Gullan and Cranston 2000, 38)
tips overlap to create a triangular pattern while the coleopteran elytra are positioned in a straight line
down the insect’s back.
In recovered fossil specimens, the remains are often disarticulated and the membranous
features are relatively rarely preserved. Regardless, because of their range of modifications and
variations, the examination of fossilised pronota, elytra, and legs can result in the identification of
insects to genus and species level.
2.2.5 The Abdomen
The typical insect abdomen is comprised of a maximum of eleven complete segments;
however, one segment is occasionally incorporated into the thorax and the last segment is usually
represented by appendages only (Gullan and Cranston 2000). Additionally, the fusing of abdominal
segments is common in many insects. Each segment is composed of two principal sclerites, a dorsal
tergum, and a ventral sternum (Borror and White 1970).
25
In most insects, the abdominal segments are void of appendages except for the posterior
eleventh segment. When the terminal appendages are present, they may include a pair of dorsolateral
cerci, a median epiproct, a pair of lateroventral paraprocts, and the genitalia (Borror and White 1970).
The cerci generally are annulated and filamentous and may be modified into feelers (e.g. mayflies) and
claspers (e.g. earwigs) (Gullan and Cranston 2000). On the eighth and ninth abdominal segments, the
genitalia are usually formed and may be internally or externally present (Gullan and Cranston 2000).
Moreover, the male genitalia are rather varied and often very complex (Borror and White 1970). As a
result, the dissection of abdominal segments towards the examination of genitalia harbours taxonomic
implications for the recognition of modern and, where preserved, fossil insects.
2.3 Identification Methods
2.3.1 Modern Insects
Keys
The employment of entomological keys permits researchers to gradually narrow down species
possibilities through the process of elimination based primarily on morphological differences of the
exoskeleton. The keys are composed of a series of couplet stages, binary opposites, which progress to
the name of the species examined, i.e. if the insect has feature A proceed to couplet 6; if not then go to
couplet 27. For example,
“1. Entire elytron uniformly pubescent, or at least with one row of setae
or bristles along entire length of each (or every second) interval.
…….2
-
Elytron glaborous, except for marginal setae and often setiferous
“dorsal” punctures on intervals 2-3, or with only outer intervals
pubescent.
…….18”
(Lindroth 1985, 24).
If the specimen matches the first alternative of couplet 1, then proceed to couplet two.
However, if the insect matches the description in the second alternative of couplet one, then continue
to couplet 18. The identification method progresses in this manner until a probable species is offered.
While the couplet method of identification is the most prevalent, a second method is available.
Rather than being confronted with dichotomous alternatives, table keys issue a single statement at
each stage which if correct the researcher advances to the next sequential statement. If the statement is
incorrect, an alternative bracketed number is listed for reference. The table style is exemplified by
Joy’s (1976) classification of Staphylinidae:
26
“1 (4). El. with strongly raised longitudinal keels.
2 (3). Segments of hind-body with 3 raised longitudinal keels on each, and ant. clubbed.
MICROPEPLINAE
3 (2). Hind-body simple, and ant. thickened to apex.
PSEUDOPSINAE
4 (1). El. without raised longitudinal keels.
5 (6). Eyes very large, and ant. clubbed characteristically.
STENINAE” (Joy 1976, 3).
While both types of keys provide a tabulated format for the recognition of insects based on
humanly delineated phenotypic structures, caution is necessary in their application because insects,
like humans, possess individual variations, resulting in a margin of error.
Thus keys are best
employed as means for quick identification of probable species. In order to more accurately verify an
insect species, the specimen in question should be compared to previously identified species from
credible entomological sources, e.g. museum collections.
Genetics
In order to compensate for the difficulties inherent in the traditional key methods, genetic
methods have been developed that provide rapid and accurate identifications (Cainé et al. 2006).
Genetic identification methods have proven proficient at species determination regardless of specimen
damage or insect life stage (e.g. Wells and Sperling 2001). However, in order to genetically determine
a species, it must have a pre-existing record in the databases, e.g. Genbank. This is established by
extracting DNA signatures from positively identified species and generating a ‘barcode’ for that
particular species (Murphy and Fraser 2006).
Processing DNA from a specimen identifies a genetic code which is unique to that species. By
noting variations in the code, different species can be recognised. For example, the partial DNA
signature, provided by the Barcode Life Project and corresponding with the mitochondrial CO1 gene,
used to identify Sitophilus granarius is:
1 ATT CTC TAC AAA CCA CAA AGA TAT CGG CAC
31 ACT ATA TTT TAT TTT TGG AGC ATG ATC AGG
61 AAT AGT TGG AAC CTC TTT AAG ACT ATT AAT
91 TCG AGC AGA ATT AGG AAA CCC CGG CTC ACT
27
121 GAT TGG AAA TGA TCA AAT TTA TAA TAC TAT
151 CGT TAC TGC TCA CGC ATT TAT TAT AAT TTT
181 TTT TAT AGT TAT ACC TAT CAT AAT TGG AGG
211 ATT CGG AAA TTG ACT AAT TCC ATT AAT ATT
241 AGG AGC CCC AGA TAT AGC CTT CCC ACG ATT
271 AAA CAA TAT GAG ATT CTG ACT ACT TCC CCC
301 ATC TTT AAT TCT TCT ATT AAT AAG AAG ATT
331 TAT TGA AAA AGG TGC TGG AAC AGG GTG AAC
(Genbank ID DQ453486);
whereas the partial code for Sitophilus oryzae is:
1 ATT CTC TAC TAA CCA CAA AGA TAT CGG AAC
31 ATT ATA CTT TAT TTT TGG AAC ATG ATC AGG
61 AAT AGT AGG TAC ATC CTT AAG TTT GCT AAT
91 TCG GGC AGA ACT AGG AAA TCC TGG ATC ACT
121 AAT TGG AAA TGA CCA AAT TTA TAA TAC TAT
151 TGT CAC AGC ACA TGC ATT CAT TAT AAT TTT
181 CTT TAT AGT AAT ACC AAT TAT AAT TGG AGG
211 ATT TGG AAA CTG ATT AAT CCC ATT AAT ATT
241 AGG AGC CCC AGA TAT AGC ATT CCC CCG TTT
271 AAA TAA TAT AAG ATT TTG ATT ACT TCC ACC
301 CTC CTT AAC TCT TTT ACT AAT AAG AAG ATT
331 TAT TG AAA AGG GAG CAG GAA CAG GATG AAC
(Genbank ID AY131099).
Sitophilus granarius and Sitophilus oryzae are both species of the same genera; however,
subtle differences are apparent in their genetic code. Although both of the gene sequences begin with
“ATT CTC TAC,” the first variation appears at the beginning of the fourth nucleotide triplet. In
Sitophilus granarius the nucleotide adenine (A) is present while the sequence for Sitophilus oryzae has
thymine (T). In these partial 360 nucleotide sequences, there are 51 base pair differences at the
genetic level between these two closely related species. By analysing these notable distinctions,
species determination is possible, and with the assistance of computer processing, the genetic
identification method is often quicker than the traditional entomological recognition methods.
2.3.2 Fossil Insects
Comparative Analysis
The study of Quaternary and late Tertiary insect remains in sediments was revitalised over
forty years ago by G. R. Coope (Elias 1994) and is based largely upon the geological principle of
28
uniformitarianism (the present is the key to the past) and the assumed relationship between the modern
ecological community, the biocoenosis, and the fossilised death assemblage, the thanatocoenosis. The
crux of this assumption centres on Coope’s argument that in most temperate and arctic environments
of the northern hemisphere, “it can be demonstrated that wherever fossils are available the Coleoptera
show a remarkable degree of morphological stability throughout the Quaternary” (1977a, 324), and
“the majority of species seems to have remained physiologically stable during this period” (1970,
107). Because of this theory of species constancy, palaeoentomologists are able to identify insect
fossils based on comparative analysis with modern species and extract palaeoecological information in
a similar fashion to anthropological analogies—superimposing the habitats of modern insects over the
fossil record.
Coope (1970) stated that the traditional elimination method of the entomological keys towards
the identification of modern species could not be utilised in the identification of fossil insect remains
based on the premise that the complete specimen were rarely recovered. Because the fragmentation of
the remains rendered the application of the keys redundant, some early entomologists erroneously
claimed that fossilised insect species could not be identified. However, the concept of an insect
species, modern and fossilised, is reliant on morphological distinctions in the exoskeleton. The
preservation of insect fossils, while often disarticulated, is such that in many cases, the heads,
thoraces, and elytra retain morphological characteristics that permit examination of detailed
microsculpture. Moreover, in contexts with excellent preservation, researchers have been able to
recover and dissect fossilised insect abdomens in which genitalia and gut contents have been preserved
(e.g. Matthews and Telka 1997). Because morphology is the primary criterion for the identification of
modern specimens, comparative analysis of modern and fossil insects can be employed for recognition
of the archaeologically recovered remains.
Although morphological characteristics are preserved, taphonomic alterations are occasionally
evident in the fossilised remains, especially when dried, and must be considered in the identification
process (Coope 1970). The original colouration of the specimens may be rendered bluish-black,
brown, or completely devoid of colour, depending on the nature of the soil in which it was retained. In
specimens in which colour remains, the colouration may have changed—red to green or green to blue.
In dried fossils, deepened puncturations may develop, and prominences and hollows may become
flattened (Coope 1970). While alterations may transpire in the post-mortem, comparative analysis of
modern and fossil insects is successful if care is given to the consideration of taphonomic afflictions.
Another taphonomically imposed hurdle in the study of fossilised insect remains is the biases
resulting from the taxonomic orders preserved, recovered, and documented. Coleoptera are the most
commonly recovered and determinable insect fossils due to the robustness of their exoskeleton and the
multitude of literature concerning the modern coleopterous fauna, which eases the identification and
interpretation process. Because of the dominance of Coleoptera in the documentary and fossil record,
29
it will contribute much of the information discussed in this work. Although coleopteran remains often
comprise the majority of recognisable fossils, dipteran (true flies) fragments are frequently recovered
though often severely disarticulated. However, researchers have demonstrated that identification of
dipteran remains can serve as an index for the reconstruction of past temperature regimes (e.g.
Skidmore 1996). The heads and propodea of Hymenoptera (sawflies, bees, wasps, and ants) are
common in archaeological assemblages and have been utilised for economic (e.g. Kenward and Hall
1995) and palaeoecological (e.g. Zazula et al. 2002) postulations. While Dermaptera (earwigs),
Hemiptera (true bugs), and Trichoptera (caddis flies) are also common, they have been studied to a
lesser extent and have not been employed to their full palaeoentomological potential. The recovery of
the membranous wings of most insects and Mallophaga (louse flies) and Anoplura (sucking lice)
specimens, which have thin flexible cuticles, is indicative of other preservational determinants, such as
soil pH, though the cuticular thickness remains the primary factor. Although fossils of Orthoptera
(cockroaches, crickets, and grasshoppers), Odonata (damselflies and dragonflies), and Lepidoptera
(butterflies and moths) are occasionally archaeologically present, their rareness in the fossil record in
comparison to the more robust fauna is not reflective of their abundance in the past.
Species constancy has permitted the identification of fossil insects through comparison with
their modern counterparts. However, archaeologically recovered populations are biased because they
provide insight into only a small point in space, which is further subjected to taphonomic distortions,
from a complex occupational site. Thus while modern analogues are applicable for identification of
fossil insects and the construction of pertinent ecological parallels, archaeological populations do not
truly resemble modern populations in regards to tabulating an accurate account of the frequency and
abundance of a species.
2.4 Ecology
Charles Elton (1927) described ecology as scientific natural history, being reminiscent of
researchers like Darwin and Linnaeus. However, the field of ecology progresses beyond the study and
taxonomic classifications of species to encompass the examination of the “total relationships
(interactions implied) of organisms with their environments, at the level of the individual
(autoecology) or that of variously constituted groups (synecology—a community ecology)” (Huffaker
et al. 1984). While ecology specialises in the analysis of ecosystems, it is an interdisciplinary
amalgamation of concepts from:
• genetics (the investigation of mutation, recombination, selection, and diversity within taxa at the
molecular level);
• physiology (the elucidation of mechanisms for cellular development and function towards the
integration of specialised tissues and organs);
30
• behaviour (the study of how the physiological components, which directly interact the external
environment, influence an individual’s reaction to and interaction in an environment);
• evolution (the evaluation of an individual’s current expression of genetic material in relation to the
various historical influences which have acted upon its species) (Krebs 1972).
An ecosystem conceptualises the interplay between the biotic and abiotic world as a whole
interlacing assemblage (cf. Tansley 1939). In this system, organisms and populations interact with
one another whilst simultaneously influencing the continuous cyclic transfer of energy and material.
Evans (1956) delineates the dynamics of an ecosystem as:
“the circulation, transformation, and accumulation of energy and matter through the
medium of living things and their activities. Photosynthesis, decomposition, herbivory,
predation, parasitism and other symbiotic activities are among the principal biological
processes responsible for the transport and storage of materials and energy, and the
interaction of the organisms engaged in these activities provide the pathways of
distribution” (Evans 1956, 1127).
The fundamental concept of archaeoentomology is founded upon this ecological
interplay having persisted in Quaternary insects unchanged. Underpinning this is the theory of
species constancy, that under repeated conditions of change, insect communities will migrate
rather than adapt through evolution (Coope 1978). Although there are studies of isolated island
and mountain populations where migration is rare and speciation dominates (Elias 1994),
extensive paleontological research has confirmed that some insects have remained
physiologically and morphologically constant for up to 30 million years (Elias 1994). Based on
this premise of stability, the ecological requirements of modern species may be analogously
employed towards reconstructions of palaeoecology.
A discussion and understanding of the major ecological principles is particularly crucial
to this study because it enables archaeoentomological researchers:
• to employ the insect fauna’s ecological associations to reconstruct environments and climates
of the past;
• to discern insect associated (directly or indirectly) materials used by past humans;
• to overlap species’ thermal requirements in order to glean an understanding of a site’s
potential maximum and minimum temperatures, which may aid in identifying species that
are ecological outliers, i.e. invasive.
2.4.1 Components
As a system, an ecosystem is composed of myriad components of various levels of complexity.
In a given locality, individual organisms comprise groups of potentially interbreeding individuals,
31
called populations (Mayr 1963).
). Species are groups of individuals in a population that are capable of
interbreeding and producing viable offspring. In turn, species may be categorised into guilds (species
that exploit the same resource by sim
similar means); guilds may form part of component communities
(species associated with some microenvironment or resource, e.g. leaf litter); and component
communities in the same area may interact to form compound communities, e.g. a pond (Price 1984).
1984
The interaction of thesee compound communities and their physical environment constitutes
constitute an
ecosystem.
Figure 2.8 Diversity of beetles in a tree habitat
(Evans 1977, 232)
In an ecosystem, the relational position of a species or population, i.e. its niche, is controlled
by habitat, resource availability, and trophic position (Elton 1927). Within a habitat [e.g. Figure 2.8],
the members of the faunal community occupy certai
certain roles—predator,
predator, herbivore, scavengers, etc. The
occupation of these roles is best evidenced in the relationship between aphids, ants, and ladybirds. On
a rose, aphids suck sap from the phloem vessels and secrete a honeydew-like residue, and ladybirds
prey upon the aphids as a food source. Additionally, ants living on the rose may try to protect the
aphids from the ladybirds in order to maintain their honeydew farm. Thus on the rose, there are
herbaceous aphids, predatory ladybirds, and mutualistic scavengers in the form of ants.
This
community is further monitored by resource availability in that if the ladybirds prey too heavily on the
aphids, there will not be sufficient aphids to feed all the ladybirds and some may die. Likewise, if the
plant dies, the aphids, which are dependent upon the rose for food, will die. While the groundwork for
these roles is omnipresent in various habitats, the animals may differ. For instance, instead of aphids,
ladybirds, and ants on a rose, there may be sheep, wolves, and shepherds in a meadow. Within
ecosystems, communities, and habitats, organisms and populations exist and interact and are able to
survive by fulfilling a specific role, which is unique to them.
2.4.2 Synecology
The concept of an ecosystem is based on the interaction of its components. Price (1984)
groups the interacting components into autotrophs, heterotrophs, and saprophages. Self-nourishing
organisms, like plants, are autotrophic in that they capable of converting and utilising light energy and
organic chemicals. Heterotrophs are organisms that feed directly or indirectly on autotrophs. If the
organisms feed directly on the autotrophs (thus transferring the energy accumulated by the autotrophs
directly into themselves), they are typically defined as herbivores, i.e. plant feeders.
However,
carnivores, which feed on the non-autotrophic organisms, still receive the autotrophic energy, though
indirectly. Additionally, saprophages, or decomposers, are considered heterotrophic because they feed
upon dead organic material; whether directly from the autotrophs or indirectly from other heterotrophs
depends on the organism. These ecological associations may be mutually beneficial or involve the
advancement of certain individuals, possibly at the expense of another.
The specific interactions evidenced by ecological associations are inter- or intraspecific as
symbiosis, herbivory, and competition (Speight et al. 1999). Symbiotic relationships may categorised
as mutualisms (a relationship in which both organisms benefit), commensalisms (an affiliation in
which one organism benefits and other remains unharmed), or parasitisms (an association in which
one species benefits at the expense of another). Myrmecophytism is a type of mutualistic ant-plant
association. During myrmecophytism, plants provide ants with food and shelter in return for defence.
Vasconcelos and Casimiro (1997) have demonstrated the myrmecophytic relationship between plants
in the genus Cecropia and Azteca alfari ants. The extrafloral nectaries of the Cecropia trees produce
glycogen which may be broken down into glucose to provide A. alfari with a food source; however,
the foliage of Cecropia is foraged by Atta laevigata, a leaf-cutting ant. Vasconselos and Casimiro
33
(1997) observed that when Cecropia housed Azteca alfari, the predacious A. alfari deterred the
foraging Atta laevigata, and the leaf-cutting ants selected other plants for harvesting. The association
between Cecropia trees and Azteca alfari mutually benefited both species, the trees received
protection and the ants acquired housing and food.
The relationship between the Ponerine ant Gnamptogenys menadensis and the Formicine ant
Polyrhachis rufipes is indicative of a commensalistic interspecific interaction which provides benefit
to one species while the other remains unaffected. Gnamptogenys menadensis foragers are reliant on
chemical trails laid down between their nests and sugar sources. Polyrhachis rufipes exploit these
trails to gain access to sugar sources. When a Polyrhachis rufipes worker encounters a Gnamptogenys
menadensis forager, the P. rufipes exhibits aggressive antennal boxing to which G. menadensis reacts
submissively (Gobin et al. 1998). By utilising the chemical trails ‘blazed’ by the Ponerine ants, the
Formine ants are able to beneficially gain access to food resources while the Ponerine ants are neither
benefited nor harmed by the association.
Unlike commensalisms, parasitic relationships aid one organism at the expense of another.
Eggleton and Belshaw (1992) approximate ten percent of all insect species are parasitic at some stage
in their life cycle. Adult females of Lysiphlebus testaceipes deposit an egg into aphids. Two days
after the egg is injected, it hatches and the larva begins to internally feed on its aphid host, resulting in
the death of the aphid six to eight days later. Upon the aphid’s death, the larva creates a hole in the
bottom of the aphid in order to attach the aphid to a leaf with silk and glue and moults to the pupal
stage. After four or five days, an adult wasp emerges from the aphid mummy (Knutson et al. 1993).
Through its relationship with the aphid, the parasitic wasp is able to conduct its life cycle; however,
the aphid does not gain any benefits and eventually dies as a result of the interaction.
Some ecological commentators have depicted herbivory as a means of parasitism (e.g. Price
1975). This theory is supported by the larvae of the European corn borer Ostrinia nubilalis, which
burrows into corn stalks and devours the fruit tissue (Gullan and Cranston 2000). By feeding on the
corn, the larvae reduce the reproductivity of the corn plant. However, it is not always the case that
phytophagy inflicts harm on the autotrophic plants, e.g. the mutualism between Azteca alfari and
Cecropia trees. Additionally, some researchers have proposed that herbivory may benefit plants.
Pruning and mowing of plants by herbivores may advance the overall reproductivity of the plant
through altering growth form and increasing quantity of seed sets (Gullan and Cranston 2000).
Unlike the specific interactions expressed by the different forms of symbiosis and herbivory,
competition is an ecological association which is indicative of a negative-negative interaction. Begon
et al. (1990) describe competition as an interaction between individuals, brought about by a shared
requirement for a resource in limited supply, and leading to a reduction in the survivorship, growth,
and/or reproduction of the competing individuals concerned. It may be intra- and interspecific.
Intraspecific competition exists between members of the same species, whereas interspecific
34
competition refers to resource competition among members of different species. At the Roman age
site of Park View School, County Durham, abundant fossil remains of the stored grain weevil,
Sitophilus granarius, were recovered (King unpublished).
During infestations of granaries with
plentiful resources, a single weevil will inhabit a single grain at any given time. However, when
resources are limited, more than one weevil will occupy the same grain simultaneously, i.e.
intraspecific competition, often resulting in reduced growth. At the Park View School site, evidence
for intraspecific competition was indicated by the presence of weevils of reduced size in addition to
specimens of normal length (pers. comm. H Kenward 2007).
While the existence of intraspecific competition is apparent, interspecific competition is less
discernable and in accordance with Gause’s Axiom (Speight et al. 1999) not possible. Gause’s Axiom
proposes that two species cannot coexist if they have identical niches as any variation in the species
may result in a competitive advantage permitting the displacement of the weaker competitor. Hence,
although Sitophilus granarius and Oryzaephilus surinamensis are both commonly recovered pests of
stored products in the same archaeological contexts, their niches may have overlapped but not been
identical. O. surinamensis is able to process meal and other ground starches but is unable to cope with
whole, dry grain unless it is damaged (Horion 1960). Therefore, O. surinamensis is not in direct
competition with S. granarius regarding undamaged grains. Rather O. surinamensis commensally
benefits from the interspecific association by processing the grains damaged by the grain weevils.
Although an overlap exists, the interspecific competition is restricted by subtle variations within the
two coexisting species niches—the presence and availability of undamaged grains provides an
unchallenged food resource for S. granarius while O. surinamensis is restricted to feeding upon the
damaged grains and scavengery. In an ideal situation, as above, resource limitation is not an issue thus
direct interspecific competition is not apparent. However, in the Park View School case, resource
limitation was proposed as an impetus for intraspecific competition amongst S. granarius. Multiple
individuals were thought to have simultaneously inhabited single grains. This would in imply that the
grain weevils were not at liberty to abandon the damaged grains for undamaged grains as resources
were limited. Hypothetically, S. granarius and O. surinamensis would have been in direct competition
for the damaged grains, and S. granarius would have had the competitive advantage by being able to
inhabit the undamaged grains first. In order to compete for the limited resources, O. surinamensis
would have had to displace or coexist with the grain weevil. The fossil evidence from a Park View
School sample tabulated an MNI of one for O. surinamensis and sixteen for S. granarius. The remains
of both species indicated little decay suggesting that cuticle robustness would not have been a strong
factor in influencing the MNI through preservation of the remains. The Park View School sample is
likely to reflect the successful competition and displacement of O. surinamensis by S. granarius
encouraged by overlapping niches and limited resources as the impetus for interspecific competition.
35
2.4.3 Ecological Constraints
Every species is limited in its distribution by biotic and abiotic factors.
In a study of
vegetational ecosystems, Holdridge and associates (1971) demonstrated that some species are
distributionally constrained to a small part of the globe by factors such as competition [see Section
2.4.2], climate, and environment, e.g. food availability. The ecological constraints that influence a
species define its niche.
Climatic factors—temperature, photoperiod, and rainfall—impinge on the ecologies of insects.
Although some species, e.g. bees and moths, are capable of elevating their body temperatures through
rapid contractions of their flight muscles, all insects are poikilotherms (their body temperature and
metabolism are functions of their surroundings) (Speight et al. 1999). Therefore, an insect’s growth,
development, and activity are dependent on temperature. For example, the saw-toothed grain beetle,
Oryzaephilus surinamensis, is restricted in its distribution in the north as it is unable to complete its
development at temperatures below 18 °C and does not flourish below 22 °C (Howe 1965).
Archaeoentomologists have employed modern species’ temperature restrictions toward
developing mechanisms for the reconstruction of palaeoclimates (e.g. Buckland 2000). If multiple
temperature restrictions are calculated for fossilised insects at a site, temperature restrictions for the
autochthonous species will overlap resulting in mutual minimum and maximum temperatures for the
site at that period in time. Using Anglo-Scandinavian York, UK as an example, the insect evidence
from 16-22 Coppergate, 6-8 Pavement, 5-7 Coppergate, and 1-9 Micklegate was utilised to calculated
the mutual climatic range [MCR] with the assistance of the BUGS Coleopteran Ecology Package
(Buckland and Buckland 2006). The MCR for each of the four sites indicated that York had a
maximum temperature of 18 °C and a minimum temperature of -7 °C during the Anglo-Scandinavian
period [see Table 4.5].
Like temperature, photoperiod, i.e. the length of daylight during a 24 hour day, is an impetus in
the development of insects. Leimar (1996) postulates that photoperiod is used by insect larvae to
ascertain information about seasonal change in order to correlate growth and development rates with
favourable conditions.
The impact of day-length is considered in Nealis and others’ (1996)
examination of the parasitic wasp, Cotesia melanoscela. When exposed to day lengths greater than 18
hours, the larval wasps developed continuously to their adult stage; however, the larvae halted
development in the cocoon prepupal stage by entering a diapause when they were subjected to less
than 16 hours of daylight. Through monitoring photoperiod, the adult parasitoid Cotesia melanoscela
arrive during the long days of summer when their hosts are most prevalent; thus improving their
chances of depositing eggs and continuing the species.
However, because of its reaction to
photoperiod, Cotesia melanoscela is restricted ecologically. If the parasitic wasp was transported to
an environment that constantly received less than 16 hours of daylight, it would never develop beyond
its larval form.
36
Rainfall can directly and indirectly affect insects. Like the insect’s relationship to photoperiod,
seasonal changes in rainfall influences species abundance. This is reflected by some species being
prolific during dry seasons and others during wet. Haggis (1996) demonstrated that outbreaks of the
cereal pest Spodoptera exempta were related to amounts of rainfall, as severe outbreaks were often
preceded by periods of drought. It was inferred that the wet conditions caused viral infections to
afflict the larvae while the heat of the droughts destroyed the viruses allowing more of the larvae to
survive to adulthood. Rainfall is also a factor in determining food availability for insects. The amount
of rainfall affects the growth of plants and as such the food source for herbivores and the species that
feed on the herbivores. In instances of drought, the plants may become stressed, which could result in
limited resource availability, increased competition, and reduced survivability for the herbivores.
The availability of food sources in an environment is a fundamental determinant in the
restriction of population size [see species competition in Section 2.4.2]. In the absence of food, an
organism is confronted with three options: relocation, diapause, or death. Through the ingestion of
food, the insects are able to obtain the energy and nutrients necessary for metabolic function, growth,
development, and reproduction (Hagen et al. 1984). When subjected to limited food availability due
to seasonal change, natural disasters, etc, insects will often relocate (through migration or short-range
dispersal) or undergo a period of facultative diapause rather than starve. Insect species which are
reliant on migration may be capable of:
• “changing behaviour to embark, such as young scale insects crawling to a leaf apex and
adopting a posture there to enhance the chances of extended aerial movement;
• being in appropriate physiological and developmental condition for the journey, as in the
flighted stage of otherwise apterous aphids;
• sensing appropriate environmental cues to depart, such as seasonal failure of the host plant
of many aphids;
• recognizing environmental cues on arrival, such as new host plant, and making controlled
departure from the current” (Gullan and Cranston 2000, 162).
An alternative to migration is diapause where the individual enters a state of arrested
development as a direct response to unfavourable conditions. Diapause may be induced and/or
terminated by environmental cues including photoperiod, temperature, food quality, moisture, pH, and
chemicals (Gullan and Cranston 2000). In the case of food availability, the period of diapause may be
ended by the reappearance of the resource. In herbivores, the diapausal termination may be instigated
by the plant’s release of chemicals or pheromones, which the insect senses.
However, one
environmental cue may override another if conditions remain unfavourable in another respect. Thus if
the food source reappears but the temperature and photoperiod remain inappropriate, the diapause will
remain in effect.
37
In section 2.4, components, habitat and trophic level were delineated as two of the definitive
measures of an ecological niche. In addition to the consideration given to where an organism lives and
its role in that habitat, the ecological constraints that act upon a species set forth the full range of
environmental conditions under which an organism can exist. In the absence of ecological constraints,
organisms would be able to develop, grow, and reproduce to abundance without limitations. However,
temperatures fluctuate, droughts occur, and food is both limited and the subject of competition. In
order to assess a species’ niche, its environmental limitations and its reaction to ecological stress must
be considered.
2.4.4 The Ecology of Invasive Species
Although, as discussed above, organisms are established in a defined ecological niche, certain
mechanisms, e.g. migration, wind or water dispersal, phoresy, and/or human transportation,
occasionally unwittingly or wittingly act upon an individual or population such that it is removed from
its original niche and supplanted in another location. If the ecological constraints of the new location
are consistent with the limitations of niche, the relocated organism(s) may, with varying levels of
difficulty, establish itself/themselves in the new location. If all other variables remain constant, the
invasive species may face competition from indigenous organisms or if available, occupy a previously
vacant niche.
Successful colonisation of new habitats requires that the first arrivals establish new viable, selfsustaining populations (Saki et al. 2001). The traits necessary for establishment vary in accordance to
habitat-type: complex and established natural community (e.g. Elton 1927), human-disturbed habitat
(e.g. Horvitz et al. 1998), and undisturbed natural island communities (e.g. Elton 1958). Saki and
associates (2001) argue that a successful invasive species will exhibit a high fecundity rate as well as
competitiveness.
The advantage of a species being able to quickly produce large numbers of offspring and outcompete competitors is apparent when confronted with an established natural community where
breeding sites are already in use, food is already being eaten, and shelters are already occupied by
other species. In order to survive, the invasive species must establish itself in a niche, often through
the displacement of one or more organisms by means of interspecific competition.
This is
demonstrated in Elton’s (1958) depiction of the introduction and spread of the Argentine ant,
Iridomyrmex humilis, in Louisiana in 1891. The species multiplied immensely, and by 1905, it had
spread throughout the southern United States and had invaded California. Smith (1935 in Elton 1958,
56) “often witnessed combats in the field between native and Argentine ants… The fact that the
Argentine ant destroys practically all the native ants as it advances makes it comparatively easy to
delimit an area infested by them.” The Argentine ant successfully invaded the southern United States
38
by rapidly reproducing and spreading and being able to out-compete the native ants for food and
space, which lends credence to Saki et al.’s (2001) hypothesis concerning traits of successful invasive
species.
However, alien species are not always faced with resistance from native species. In areas that
have been modified or destroyed by human influence or natural disasters, niches may be vacant. The
modification or destruction of an area may temporarily ‘empty’ previously occupied niches by
displacing or killing the organisms who resided in them. Moreover, major alterations to a habitat may
result in the indigenous organisms being no longer suitably adapted to the area. When the European
colonists arrived in the New World, they cleared forest and brush for the construction of settlements
and the cultivation of crops. This land clearance displaced some of the indigenous species and created
vacant niches for the invasive species. Lindroth (1957) tabulated 638 species of insects common to
Europe and North America, and of these, he estimated that 242 were accidentally introduced through
probable association with the materials employed as ship ballast. The European insect species, which
inhabited the rubble and soil used as ship’s ballast, would have been able to carve out a niche in the
disturbed land around the colonial settlements and have faced little competition from the indigenous
species which were displaced by the human-induced modifications. The successful establishment of
Old World species in North America helped create what Crosby (2004) called Neo-European
ecological spaces, which he proposed played a key role in early settlement viability.
Elton (1958) has demonstrated the susceptibility of remote islands to invasive species. The
early ecological surveys of Easter Island detected only five endemic invertebrate species: a land snail,
a water beetle, a weevil, a fly, and a green lacewing; in comparison to the 44 types of invertebrates, 2
lizards, 2 species of bird, and rats that are known to have been introduced (Elton 1958). Because of
their remoteness, island populations are isolated. Unlike continents where several dense populations
are in constant interaction, the ‘native’ fauna of remote islands is likely to be composed of a few
species that arrived haphazardly over time from the nearest land masses, possibly through wind or
water dispersal. As a result, competition for resources and space is minimal and vacant niches exist.
The only conflicts with which an invasive species is confronted concern the ecological constraints
inherent to that species.
The ecology of invasive species and population dynamics are major components in the present
investigation.
In order to elucidate culture contact through utilisation of insect fossils, the
archaeological contexts are examined for evidence of foreign species.
However, do ecological,
morphological, and genetic variations between the native and foreign organisms exist, and are they
visible in the fossil record?
39
Chapter 3
Methodological Review
40
3.1 Introduction
Building upon the entomological principles discussed in the previous chapter, Chapter 3 will
outline the specific analytical methods of the study. The means for data collection, processing, and
recording will be reviewed, and each of the four research approaches will be detailed. First, the
palaeoecological method, incorporating habitat association and Mutual Climatic Range, is
summarised. This is followed by an outline of the biogeographical approach for assessing species
movement and the importance of absence. A review of the isotopic method comes next. The chapter
is concluded with a section discussing genetic applications and phylogeography.
3.2 Data Collection
As already noted, the aim of this investigation is to explore various methods of using insect
fossils to help trace human migration and trade. This book will diachronically and synchronically
focus on material from Neolithic and Roman England and Europe. An Anglo-Scandinavian site will
also be considered in Chapter 4. Thus, the study comprises the following components:
• A survey of the published insect remains from Mediterranean sites broadly dated to the Neolithic
and Roman periods;
• Application of palaeoecology and modern analogues to identify specific human transported species
beyond their natural geographic ranges;
• Comparison of British and European material to facilitate the recognition of patterning, and thus
allow one to infer the effects of trade, migration, and acculturation.
3.3 Processing Methods
Although the early methods, prior to the mid-1960s, of processing samples for insect remains
involved splitting sediments along the bedding planes and searching the exposed areas for fossils
(Coope 1959), the method proved to be time consuming and had a tendency for bias as it led to the
more conspicuous insect remains being over represented. In 1968, Coope and Osborne (cf. Coope
1986a) presented an alternative procedure based on a wet sieving method. Once their technique was
modified to include a flotation stage designed to concentrate the cuticle, it has since proven effective.
While the paraffin flotation method is both cheap and efficient, it is not without problems and several
amendments aimed at increasing fossil recovery have been presented (Kenward 1974; et al. 1980;
1985). However, these modifications differ only slightly from the original methodology employed by
Coope and Osborne (1968). Rousseau (2009) has systematically tested the paraffin approach and
provides a detailed discussion of its efficiency, or rather its inefficiency.
41
3.4 Palaeoecology
3.4.1 Theoretical Perspective
As discussed in Chapter 2, organisms are established in a defined ecological niche which in
turn is regulated by habitat, resource availability, and trophic position (Elton 1927).
When an
organism is transported to a foreign location, its survivability is dependent upon factors such as
ecological constraints and competition. In studying the past, palaeoentomologists have relied on the
analysis of modern insect species which firstly, produces ecological data on specific species and
species groups, and secondly, can be used as controls to be contrasted with the fossil species
associations. Additionally, the work of Kenward (1975a; 1976; 1985) on the ‘background’ component
of modern death assemblages has served to emphasise the need for caution in interpreting fossil insect
assemblages. Through modern analogues, palaeo-entomologists are able to ascertain information
about ecological components, species associations, and behavioural characteristics of various species
and apply the knowledge to the fossil remains.
3.4.2 Background to the Methodology
Ecological Categorisation
In order to assess the potential of archaeological insect assemblages in the reconstruction of
products and exchange patterns, invertebrate remains must have been retrieved from the sites and their
associated ecology delineated. By assigning the insect fauna to broad ecological categories, it is
possible to separate the autochthonous species from those individuals that may have been transported
on materials from other habitats (Kenward 1974). The broad grouping method can be problematic
with insects because of regional variations in species, differences in ecological niches based on the
life-cycle stages of a single species, and habitat seasonality (Hill 1994b). While Hill (1994b) cautions
about the disadvantages of using broad ecological classifications, it is applied in this study to help
identify the indigenous species that would not have been imported.
Once the autochthonous
individuals have been recognised, it will be possible to examine the remaining insect species for
associations with specific products and habitats, which could be utilised to detect exchange patterns.
A variety of schemes exist concerning the ecological grouping of archaeologically
recovered insect remains. Kenward (1978ab) and Hall et al. (1983) employed an ecological grouping
system based on a limited range of categories. Kenward (1978a) emphasised the importance of coding
for outdoor species and aquatic and aquatic-marginal fauna, and Kenward (1979) incorporated
categories of species exploiting decomposing matter. Hall and Kenward (1980) and Hall et al. (1983)
formalised the system through its use in categorising a large insect assemblage. Kenward (1988) and
Hall and Kenward (1990) added further ecological groups including wood and bark, with stored grain,
42
with living plants, and with heathland/moorland. Furthermore, a synanthrope group was incorporated
and utilised by Kenward (1997).
Robinson (1981; 1983) designated ten species ecological groupings: aquatic, pasture/dung,
probable meadowland, wood and trees, marsh/aquatic plants, bare ground/arable, dung/foul organic
material, Lathridiidae, synanthropes, and species especially associated with structural timbers. Later,
an eleventh group was added, i.e. species on roots in grassland (cf. Robinson 1991). A similar system
was employed by Hellqvist (1999), and Hill (1993) produced a classification closely modelled upon
Robinson’s.
Hill (1993) specified the following groups: eurytopic, aquatic, synathropic/urban,
arable/disturbed ground, pasture/dung, marsh/fen, heath/moorland, decomposers/litter dwellers,
associated with trees, true woodland species, and uncoded. The true woodland species category was
further subdivided into predators, deadwood (saproxylic), phytophagous, bark beetles, fungus feeders,
litter dwellers, ant associates and dung beetles.
A number of other classification systems have been employed by palaeoentomological
researchers (e.g. Girling 1980; Hakbilj 1989; Smith 1996ab), which does not allow for the existence of
a unified, common, ecological coding system in the discipline at this date. While a number of the
classification systems have their merits, they tend to be designed to meet the needs and the whims of
their users.
For example, Bain (1998) employed a limited number of high-level groups: pests,
compost and dung dwellers, carrion beetles, and mould and fungus feeders; whereas, Boswijk and
Whitehouse (2002) used an ecological coding classification that subdivided the species groups into a
finer level of detail than any of the systems discussed above. The narrower the resolution, the more
equipped the system is at detecting subtle environmental changes within and between contexts.
Palaeoclimatic Reconstruction
Fossil beetle assemblages have been used for more than 25 years towards reconstructing past
temperature conditions (e.g. Coope 1977a; Atkinson et al. 1987; Elias 1994). The Mutual Climatic
Range (MCR) method is one of the leading methods through which these reconstructions are carried
out.
Under this method, reconstructing the climatic conditions associated with a fossil beetle
assemblage involves three steps:
1. Modern distributional and climatic data are utilized to measure the climatic ranges or
envelopes of the species present in the fossil assemblage. These climate envelopes are usually
two dimensional; one dimension is the mean temperature of the warmest month (TMAX) and
the other is either the mean temperature of the coldest month (TMIN) or the difference
between TMAX and TMIN (TRANGE);
2.
The climatic conditions are determined using the overlap of these climatic envelopes where
the overlap itself contains a range of climatic conditions;
43
3. The range is issued single values of TMAX and TMIN (or TRANGE). This conversion is
founded on linear regression models relating observed modern values of these variables to the
corresponding midpoints of the ranges found through application of the first two steps to the
modern data.
3.4.3 Methodology
The palaeoecological aspect of this work is two-fold; firstly modern analogues will be used to
tabulate the behaviour, habitat, and associations of the fossil entomofauna in order to reconstruct the
palaeoenvironment of a site, and secondly, the MCR method will be employed to estimate the
palaeotemperature of a site [Chapter 4]. The utility of any individual species which is intended to be
employed as a palaeoecological indicator is reliant upon the ecological parameters which limit its
distribution in time and space. Consequently, stenotopic insects are of greater value than eurytopes as
they have narrower habitat restrictions.
Because of their narrow habitat restrictions, certain stenotopic insects can be used as indicators
of the presence of associated material or conditions in the environment. Moreover, as this study is
concerned with human activity, the number of stenotopic fauna assessed can be further reduced with
the consideration of only synanthropic insects (species which are unable to maintain breeding
populations without the ameliorated conditions provided by human buildings and/or human activity).
On this basis, the fossil material will be surveyed to identify insect species that are strongly associated
with potentially tradable commodities such as cereals. Once the potential trade indicator species have
been highlighted, MCRs will be calculated for the corresponding sites as well as a selected range of
similarly dated sites.
The TMax and TMin for each site will be compared to the temperature
requirements, especially the optimal range, necessary for the associated insect species to complete
their development. It is hypothesised that insect species whose temperature requirements fall outside
the MCR determined temperature range for a site will represent hitchhikers introduced to the site
through the importation of goods. While these species may have been able to survive within the
artificial microhabitats of human structures, if the conditions in the surrounding natural environment
presented ecological constraints on their development, it is unlikely that they would have been natives.
3.5 Biogeography
3.5.1 A Brief Literature Review
As the study of living things in space and time, biogeography addresses issues such as the
distribution of species throughout time, the mechanisms behind the distribution, and the human
influence upon these patterns of species distribution (Cox and Moore 2000). To a limited degree,
some previous applications of biogeographical concepts and palaeoentomology have been attempted.
44
The bulk of these studies have been concerned with the context of North Atlantic faunal connections
(e.g. Buckland 1988; Buckland et al. 1995; Coope 1986b; Sadler 1991a; 1991b; Sadler and Skidmore
1995; Kenward 1997). However, while all these investigations emphasise the benefits of utilising
biogeography, they restrict its use to a single site or region, note probable invasive species, and as a
result, offer tentative speculations about their origins based on historical documentary accounts.
Although these accounts have primarily hypothesised about the origins of the invasive species, they
were proficient at noting the presence and absence of species through the different chronological
periods of the site or region.
Kenward (1997) attributes the limitations of the field to a paucity of securely dated material
and palaeoentomological evidence, particularly outside of the United Kingdom; however, efforts have
been made to apply biogeography on a larger scale (to a limited extent Buckland 1981, and to a larger
degree Buckland and Sadler 1989; Kislev et al. 2004). While palaeoentomological surveys may not
have been conducted extensively outside the United Kingdom, an investigation of both the published
and grey literature on the subject permits the mapping of species’ distribution and movement through
time on a broader scale.
3.5.2 Approaching the Problem
Before biogeographical analysis can begin in earnest, a number of preliminary tasks need to be
undertaken. As the investigation is ultimately intended to address the issue of human movement and
trade in the past, it will not be realistic to attempt to apply the method to every insect species. Rather
the project relies upon the ability to target synanthropic species associated with human transported
materials.
Once the applicable insect species have been selected, an exploration of the literature will
ensue in order to document the presence and absence of the species temporally and geographically
[Chapter 5]. This information will be recorded and assessed for each species. As the existing
documented material presumably represents only a fraction of the original biocoenosis in the past and
does not account for the thanatocoenosis of future sites, it must be understood that the biogeographical
mapping of palaeoentomological remains only provides a provisional reflection of the past. With that
in mind, the resulting biogeographical interpretation will indicate the geographic presence of each
species at archaeologically dated points in time. As the species are strongly synanthropic, their
movement will hypothetically reflect human movement rather than geographic range shifts resulting
from climate change or seasonal migration.
45
3.6 Isotopic Analyses
3.6.1 History
In recent years, isotopes, especially deuterium and oxygen, have proven beneficial in
paleotemperature estimates especially from deep-sea sediments (e.g. Shackleton and Opdyke 1973;
Bauch and Erlenkeuser 2003), ice cores (e.g. Dansgaard et al. 1989; Grootes et al. 2002), and calcium
carbonate rich lake sediments (e.g. Oeggl and Eicher 1989; Talbot 1990). Given the success of
isotopes in these past studies and the application of beetles in climate reconstructions using MCR,
researchers (e.g. Gröcke et al. 2006; Hardenbroek 2006; et al. 2007) have sought to use stable isotopes
from beetle chitin to ascertain palaeoclimatic information.
Stable-isotopes have also been employed towards the tracing of animal and human diet. This
is particularly evidenced in studies investigating marine versus terrestrial foodwebs. The stableisotope composition of marine biomes tends to be more enriched relative to terrestrial C3 or freshwater
foodwebs (Peterson and Fry 1987; Schaffner and Swart 1991; Hobson et al. 1997; Hebert et al. 2009).
Carbon-13, nitrogen-15, and sulfur-34 are especially evidenced by this pattern and are commonly
analysed; however, deuterium and oxygen-18 have also shown enrichment in marine systems
compared to terrestrial (Fry and Sherr 1984; Schaffner and Swart 1991). In terrestrial ecosystems,
analysis of stable-isotopic carbon provides insight into plant resources within a foodweb. Differences
in plant photosynthetic pathways result in distinct isotopic differences between C3, C4, and
Crassulacean acid metabolism (CAM) plants (Peterson and Fry 1987; Tieszen and Boutton 1988).
Moreover, isotopic analyses may prove invaluable as indicators of geographic origin. Hobson
and associates (1999) have used deuterium and carbon-13 to trace the migration of the Monarch
butterflies to their natal origins through examination of variation in continental gradient evidenced
through the deuterium, and carbon-13 and nitrogen-15 have been shown to be linked to altitude and
humidity (Körner and Diemer 1987; Körner et al. 1988; Körner et al. 1991). Migratory connectivity
over a large geospatial scale has been investigated using stable carbon, sulfur and hydrogen for
migrant birds and insects (Chamberlain et al. 1997; Hobson and Wassenaar 1997; Wassenaar and
Hobson 1998; Rubstein and Hobson 2004). Recently, applications have been developed to assist in
systematically determining the origins of isotopes through the creation of basemaps from the
interpolation of precipitation isotope values, and the methods been employed towards studies in
wildlife forensics (Bowen et al. 2005).
46
3.6.2 Theoretical Basis
The stable isotope deuterium is a rare
component of the water molecule (H2O) and enters
organisms through means of the hydrological cycle
[Figure 3.1; Figure 3.2].
The deuterium/hydrogen
(D/H) ratio of meteoric water is a function of
temperature or climate (Dansgaard 1964), and varies
with respect to geographical parameters such as
latitude, altitude, continentality, and intensity of
precipitation as well as ecological parameters such as
trophic level. The geographical variation is the result
of the Raleigh distillation, which reflects changes in the
degree of rain-out of moisture from an air mass
(Yurtsever and Gat 1981). The water vapour that forms
Figure 3.1 The hydrological cycle
(National Weather Service n.d.)
precipitation is depleted in heavy isotopes, such as
Figure 3.2 Isotopes in hydrological cycle
(IAEA 2000)
deuterium, relative to ocean water. As condensation forms raindrops from a cloud, the heavy isotopes
condensate first, creating isotopically enriched rain, as the cloud becomes isotopically depleted by
rain-out (IAEA 2000).
This means that successive rainfall events from an air mass become
isotopically lighter. At middle and high latitudes, isotopes are closely correlated with temperatures,
and thus seasonality, with winter precipitation being more enriched than summer precipitation. At
lower latitudes, the isotopic content is a function of the volume of the precipitation. During periods of
increased precipitation, the isotopes are more depleted in the precipitation. The intensity of the
precipitation is also a factor as lighter rains are more enriched than harder rains (IAEA 2000).
The isotopic variation apparent in a region’s meteoric water is transferred to organisms living
within that region via the food web. The relationship of the D/H ratio of chitin and the D/H signature
of the meteoric water in an insect’s habitat has been documented (e.g. ladybirds in Ostrom et al. 1997,
and Monarch butterflies in Wassenaar and Hobson 1998). The stable isotopic signature contained in a
terrestrial insect’s chitin is dependent upon its diet during the formation of its exoskeleton. Thus the
hydrogen isotopes present in the chitin of predatory beetles will be based upon the isotopic values of
their prey, and herbivorous insects will reflect the corresponding isotopic signature present in the
vegetation upon which they feed. As such, a region’s isotopic indicator is transferred from the
meteoric water to the locally grown vegetation to the
herbivores to the predators.
A similar process occurs in regards to stableisotopic carbon and nitrogen. Carbon-13 is transferred
to organisms through the carbon cycle [Figure 3.3],
and nitrogen-15 is introduced via the nitrogen cycle
[Figure 3.4]. As with deuterium, carbon-13 and nitrogen-15 vary in relation to geogr-aphic and ecological
parameters. The insect species consuming food from a
specific locality will acquire the carbon and nitrogen
signature of that food and that region.
However,
fractionation occurs within the carbon and nitrogen
isotopic ratios relative to trophic level. For example,
the isotopic values are enriched in carnivores relative
to herbivores within a food web.
Figure 3.3 The carbon cycle
(Gardiner 2008)
The potential of isotopes as geographic indicators is significant. If a locally grown plant is
transferred outside of its original region through means such as trade, then herbivores feeding upon it
at other destinations will display the isotopic signature of the plant’s host region. Similarly, predators
feeding on those herbivores will carry that plant’s isotope value. By comparing the stable isotopes of
beetles that are known to feed upon local vegetation (e.g. trees and grasses) and beetles suspected of
feeding on potentially non-local plants (e.g. cereals), it is hypothesised that imported materials can be
identified.
Figure 3.4 The nitrogen cycle
(Wojciechowski and Mahn 2006)
3.6.3 Methodology and Objectives
In order to explore the potential of beetles as isotopic indicators of trade, modern and fossil
material were analysed. The modern aspect of the experiment was two-fold; involving laboratoryreared granary weevils [Chapter 6] and beetles retrieved from traps in the wild [Chapter 7]. The
laboratory experiments used a parent population (G1) of the granary weevil Sitophilus granarius L. as
a control. Ten individuals of G1 were placed in tubes containing separate cereals and seeds from
various regions with known isotopic records. In order to breed and lay eggs, the granary weevils were
left in the tubes for ten days at 28 °C. Afterwards, G1 was removed from the tubes and individuals
were held in the freezer for isotopic analysis. After 3-5 weeks, the second generation adults (G2)
emerged from the cereals and seeds in the tubes. At this stage, two G2 individuals from each tube
were frozen for isotopic analysis (G2a) and five G2 individuals from each culture were placed in
separate containers with grain from another isotopic region (G2b). At 10, 15, 20, 25, and 30 days, an
individual from G2b was removed from the containers and frozen for evaluation. It is hypothesised
that the G2 specimens will reveal the respective isotopic signature of the cereals in which their
exoskeleton was formed and not that of the G1 generation. Additionally, it is suspected that the
transfer of G2b to another cereal source will not impact its isotopic value, and that the G2b specimens
will not differ from the corresponding individuals, G2a, from their original tubes.
Similarly, ten individuals of G1 were placed in tubes containing a mixture of 25% barley, 25%
wheat, 25% oats, and 25% buckwheat kernels. The G1 specimens were allotted ten days at 28 °C to
oviposition and were then removed from the tubes. As the G2 individuals emerged after 3-5 weeks at
28 °C, they were frozen for isotopic evaluation [see Chapter 6.3.2].
Pitfall traps were employed to collect modern beetle species from the reconstructed AngloSaxon village located at West Stow, UK. The wax (lipids) and proteins were removed from the
remains; thus isolating the chitin. δ2H, δ13C and δ15N analyses were performed on the chitin. The
stable-isotope ratios were compared between the species and trophic level variations considered. The
West Stow specimens were investigated for recognition and classification of species considered to
represent local and imported materials.
In regards to the fossil material [Chapter 7], recovered archaeological insect remains were
investigated from the Neolithic sites of Eythra, Plaußig, and Erkelenz-Kückhoven, Germany.
Deuterium, carbon-13, and nitrogen-15 isotopic assays were conducted on the chitin isolated from the
individual species. The resulting isotopic predictions were compared at intra- and inter-site levels to
determine the presence of patterns in results and to identify isotopic outlier species, which may
indicate human activity.
The objective of incorporating isotopic analyses into this assessment of trade and migration is
to devise a way of differentiating between potentially tradeable perishable goods from those locally
supplied, and secondly to begin to trace the origins of tradeable products through the mapping of their
inherent stable isotope values of their respective regions in the past.
3.7 Phylogeography
3.7.1 The History of Ancient DNA
In 1984, Russ Higuchi and associates published a revolutionary finding—that traces of
deoxyribonucleic acid (DNA) from a museum specimen of the Quagga, Equus quagga, (an Equid
believed to have gone extinct in the late 19th century) remained in the specimen over 150 years after
the death of the individual and that the DNA could be extracted and sequenced (Higuchi et al. 1984).
Svante Pääbo expanded upon Higuchi’s discovery and demonstrated that the procedure could be
replicated in mummified human samples dating back several thousand years (Pääbo 1985a; 1985b;
1986).
However, until the development of the Polymerase Chain Reaction (PCR) (Mullis and Faloona
1987; Saiki et al. 1988) in the late 1980s, the field of ancient DNA (aDNA) progressed slowly. The
introduction of PCR heralded a series of papers claiming authentic DNA could be extracted from
specimens that were millions of years old. The majority of the claims were founded on the retrieval of
DNA from amber-preserved specimens. Preserved insect DNA was reported from amber-encased
fossils dating to the Ogliocene (25-35 mya), e.g. stingless bees (Cano et al. 1992a; Cano et al. 1992b),
termites (DeSalle et al. 1992; DeSalle et al. 1993), and wood gnats (DeSalle and Grimaldi 1994), and
the Cretaceous (120-135 mya) periods, e.g. Lebanese weevils (Cano et al. 1993).
50
Moreover, aDNA retrieval was not limited to amber. Golenberg and collaborators (1990;
Golenberg 1991) extracted DNA from sediment-preserved plant remains dating to the Miocene.
Additionally, DNA sequences were investigated from dinosaur bone, which dated over 80 million
years ago (mya) (Woodward et al. 1994), and Cretaceous egg (An et al. 1995; Li et al. 1995). These
sequences of DNA stretching millions of years into the past were referred to as Antediluvian DNA
(Lindahl 1993b).
Unfortunately, despite these exciting and potentially revolutionary claims, a critical review of
ancient DNA literature indicates that few recent studies have succeeded in amplifying DNA from
remains older than several hundred thousand years (cf. Willerslev et al. 2003). Since the early studies
may reflect the amplification of contaminated material, they should be regarded carefully. However,
recent investigations of sub-glacial deposits in Greenland (Willerslev et al. 2007) and insect carapaces
from museum collection samples (less than one hundred years old, e.g. Zakharov et al. 2000;
Junquiera et al. 2002; Gilbert et al. 2007) have yielded DNA fragments with no indication of
contamination.
3.7.2 The Extraction of Ancient DNA from Samples
Although the methods utilised to extract DNA from ancient specimens vary in accordance to
the tissue, the majority of aDNA studies employ one of two procedures. These techniques call for an
initial digestion of the tissue to release DNA followed by a purification step involving either organic
solvents or the DNA binding properties of silica. The experiments carried out for the purpose of this
study will rely on the silica method for purification.
To remove any surface contaminants, a specimen is typically ‘pre-prepared’ post-sampling
using a range of decontamination techniques. In non-destructive methods (as employed in this study),
the material is not ground. Instead the fossil and modern specimens are placed in tubes, fully
immersed in digestion buffer (400 µl), and incubated overnight at 55 °C with gentle agitation. As in
Gilbert et al. (2007), the digestion buffer is modified from Pfeiffer et al. (2004) to consist of 5 mM
CaCl2, 1 % sodium dodecyl sulphate (SDS), 40 mM dithiotreitol (DTT), 2.5 mM ethylenediamine
tetraacetic acid (EDTA), 250 mg/ml proteinase K, 10 mM Tris buffer pH 8, and 10 mM NaCl (final
concentrations). After incubation at modest temperatures (50-60 ºC) with gentle agitation for 16-20
hours, the nucleic acids are purified from the solution.
Silica-based DNA extractions
DNA extractions involving silica (Boom et al. 1990; Höss and Pääbo 1993) require the
extraction of DNA in high concentrations of salts such as guanidinium thiocyanate (GuSCN). The
salts have the ability to lyse proteins and simultaneously act as a chaotropic agent facilitating the
binding of DNA to silica particles.
51
After the period of incubation and gentle agitation, 2 ml of Phosphate Buffer (PB) buffer (the
protein-lysing salt) are added to the 400 µl digestion buffer and vortexed gently. Once mixed, 650 µl
aliquots are added to the spin column, spun at 6000 g for 1 minute. Then collection tube is emptied.
This step is repeated three times to ensure that the extraneous non-DNA material has been separated
and removed and that the remaining DNA has sufficiently bound to the silica filter. 500 µl PE buffer
(containing ethanol to wash the filter) is added to the column and spun for 1 minute at 10000 g in
order to rinse the filter. Afterwards, the collection tube is emptied and the column spun again at
maximum speed for 3 minutes to dry the filter. The filter is then transferred to a clean new 1.5 ml
Eppendorf Biopur tube and 50 µl of AE Elution buffer is placed on the filter and left for 5 minutes to
initiate the release of the DNA from the silica filter. Finally, the column is spun again at >10000 g for
1 minute to allow the solution to migrate to the base of the new tube.
Analyses incorporating PCR
PCR is a reiterative process that depends on the annealing of sequence specific oligonucleotide
probes (‘primers’) to complementary DNA sequences. Two primers per reaction are typically used,
each approximately 20 base pairs (bp) in length and designed to bind to the 5’ end of the target
sequence. Deoxyribonucleotide bases (dNTPs), a DNA polymerase enzyme (e.g. HiFi), 10x PCR
buffer, and 2.5 mM MgCl2 are added to a mixture of DNA, primers, and bovine serum albumin (BSA).
The specific DNA sequence can be exponentially amplified through a cyclical process involving the
repeated denaturation of templates, the binding of primers to DNA targets (annealing), and strand
replication by the enzymes (elongation).
A PCR, in theory, should result in the sole amplification of the target regions of DNA specified
by the selected primers. However, because of the likelihood of sequence modification resulting from
post mortem DNA damage and contamination, PCR reactions occasionally amplify multiple DNA
sequences, which can result in unreadable sequence data or conflicting base calls. By cloning the PCR
products, it is possible to avoid these problems.
The molecular cloning of ancient DNA
Through molecular cloning techniques, sequence heterogeneity can be detected within a single
PCR reaction. Amplicons are inserted into bacterial plasmids, which are transformed into Escherichia
coli cells.
The resulting colonies can be identified through blue/white screening as a result of
disruption to one of a range of subunits of the plasmid’s b-galactosidase gene, which metabolises Xgal (5-bromo-4-chloro-3-indolyl-b-D-galactopyranoside) into a blue product. Because each clone will
have incorporated a single amplicon, post-mortem DNA damage, jumping, and contamination can be
assessed by screening multiple clones.
52
3.7.4 Ancient DNA Authentication Criteria
Because of the risk for contamination and the occasional lack of result reproducibility
prevalent in the early Antediluvian DNA investigations, a list of suggested criteria has been published
to help limit the effect of sample contamination. Based on the criteria proposed for forensic studies
(Carracedo et al. 2000), this list is designed to help minimise some of the field’s errors (Handt et al.
1994; Cooper and Poinar 2000). The list is as follows:
1) Isolation of work areas in an effort to avoid contamination by separating the environments used for
the handling of samples and extraction of DNA from the PCR amplified products;
2) Negative control extractions and amplifications to monitor for contaminants infiltrating the process
at any stage;
3) Appropriate molecular behaviour – because of DNA degradation, aDNA investigations should be
suspicious of the successful amplification of large DNA fragments;
4) Reproducibility – multiple PCR and extractions should generate consistent results;
5) Cloning of products to screen for damage, contamination and jumping PCR;
6) Independent replication – the generation of consistent results by independent research groups;
7) Biochemical preservation – preservation of other biomolecules that correlate with
DNA survival (e.g. collagen, amino acid racemisation) should be indicative of good sample
preservation;
8) Quantification - through competitive PCR or real-time PCR to give an estimate of the number of
starting templates in the reaction;
9) Associated remains – are associated remains equally well preserved, and do they show indicate
contamination?
3.7.5 Targeting Mitochondrial DNA: The Aims of the Project
Mitochondria are endosymbiotic organelle that resemble free-living eubacterium similar to
modern α-Proteobacteria (Gray et al. 1999; Lang et al. 1999) and act as a cell’s respiratory source for
the generation of ATP. There are thousands of mitochondria in each cell, and its presence in copy
numbers is approximated to range between 1,000-10,000 times that of single-copy nuclear DNA
(Taanman 1999). As a result, given similar rates of degradation, mitochondrial DNA will remain
present in a cell longer than nuclear DNA.
In addition to this, mitochondrial DNA (mtDNA)
undergoes a fairly rapid rate of evolution (Lang et al. 1999) and is inherited maternally which makes it
exceedingly useful in population studies.
Animal mitochondria span 16-20 thousand base pairs in length, containing the same 37 genes
coding for small- and large-subunit rRNAs, 13 proteins, and 22 tRNAs (Hoy 2004) and at least one
53
noncoding (control) region (Boore and Brown 1998). In insects, this noncoding region is rich in
adenine (A) and thymine (T) and regulates replication and transcription (Hoy 2003). In modern
phylogeographic studies, researchers have primarily focused on the cytochrome oxidase I (COI)
(specifically the variable portion corresponding with sites 2410 and 2665 in the Drosophila yakuba
mitochondrial genome) and cytochrome oxidase II genes (e.g. Juan et al. 1998, Moya et al. 2004;
Smith and Farrell 2005).
Given the suitability of the mitochondrial genome for phylogenetic investigations, the present
study aims to investigate the retrievability of genetic information from archaeologically recovered
insect remains of the cereal pest beetle Sitophilus granarius L. from waterlogged contexts dated to the
Roman and medieval periods and seeks to investigate the variation presented in the intraspecific
mitogenomes towards the potential mapping of geographic relationships between populations of
various trade-indicator species past and present. This initial investigation will involve the application
of PCRs in order to amplify key regions of the COI and COII genes in order to pinpoint variation
within species.
By conducting population studies on trade-related species, this investigation hypothesises that
following concepts can be assessed:
• The mapping of past trade patterns through a genetic evaluation of population movement and
relationships;
• The conceptualisation of the intensity of trade—the presence of high or low initial genetic
diversity;
• The trade continuity—in the case of limited genetic diversity, the presence or absence of
evidence of founder effect possibly resulting from maintained trade with a singular source
over time.
54
Chapter 4
The Palaeoecological Approach:
An Assessment of Two Case Studies
55
4.1 Introduction
In Chapter 4, palaeoecological reconstructions are formulated from the insect fossils recovered
from two urban archaeological sites that were excavated in the city centre of York, UK: 7-15
Spurriergate and 16-22 Coppergate.
The palaeoenvironmental reconstructions roughly follow
Robinson’s (1981; 1983) ecological coding system. The following species groups are employed:
Group 1: Aquatic
This category includes all beetle species that can spend most of their adult life under water,
e.g. Helophorus sp.;
Group 2: Pasture/dung
Species Group 2 is composed of dung beetles which mostly occur in or under the dung of large
herbivores. The species are more common with dung in the field than manure heaps. It is
usually comprised of species from the genera Geotrupes, Copris, Aphodius, and Onthophagus;
Group 3: Probable meadowland
This category is comprised of species which mostly feed on leaves and stems of vetches,
clovers, and other grassland flora, e.g. Mecinus pyraster and Sitona sp.;
Group 4: Wood and trees
Beetles which are found in the wood, leaves, bark, and fruits of live trees and shrubs as well as
species which feed on wood that is undergoing various stages of decay, e.g. Magdalis
carbonaria;
Group 5: Marsh/aquatic plants
These are species of beetles which feed exclusively on marsh or aquatic plants, e.g. Notaris
acridulus;
Group 6: Disturbed ground/arable
This category includes Coleoptera that inhabit bare ground, arable soils, and weedy disturbed
ground, e.g. Amara sp.;
Group 7: Dung/foul organic material
This group consists of species which live in different types of foul organic matter such as
decaying vegetation, dung, compost, carrion, and manure heaps. The beetles are primarily
decomposers, e.g. Megasternum obscurum and Cercyon sp.;
Group 8: Lathridiidae
This classification comprises a family of beetles which primarily feed on fungi and mould on
decaying plant material, e.g. Lathridius minutus group;
Group 9: Synanthropes
56
This category consists of species which are associated with human-made environments. It is
comprised of species that usually inhabit or are associated with human-made structures, e.g.
Ptinus fur and Typhaea stercorea;
Group 10: Species especially associated with structural timbers
Coleopteran species, such as Anobium punctatum and Lyctus linearis, which live in dry dead
wood and are able to reproduce in structural timbers, are categorised in this species group; and,
Group 11: On roots in grassland
This species group includes Scarabaeidae and Elateridae which as larvae feed on the roots of
grassland herbs, e.g. Hoplia philanthus and Phyllopertha horticola.
The palaeoenvironmental reconstructions serve to provide insight into the flora, fauna, and
geography in the vicinity of the site.
By examining the habitat and diet of the insect fauna, it is also
possible to identify indicator species of commodities, which may have been exploited by humans in
the past. Several of these commodities were not likely to have been autochthonous to the sites and
may indicate palaeoeconomic activities such as long-distance trade or exploitation of the hinterland
resources.
Mutual Climatic Reconstruction models (MCRs) are used to build palaeoclimatic
reconstructions. The MCRs are compared to the thermal requirements of individual insect species to
help identify ecological outliers, i.e. species that may not have been indigenous to or capable of
surviving in the wild of the United Kingdom. The application of MCRs in conjunction with specific
‘economic’ indicator taxa is hypothesised to aid in the ability to recognise the occurrence of local and
long-distance trade.
4.2 Case Study 1: 7-15 Spurriergate, York (site code: 2000:584)
4.2.1 Introduction
Biological samples were collected during archaeological work on 7-15 Spurriergate, York by
MAP Archaeological Consultancy Ltd. in 2000 and 2005. The deposits were of early Roman (1st
century AD) to medieval date with preservation ranging from good to exceptional (cf. Hall et al. 2000;
Carrot et al. 2005). Figure 4.1 shows the location of the site to the northeast of the River Ouse near
Micklegate Bridge.
4.2.2 Processing Methods
Six samples were selected from the early Roman context 6063 for palaeoecological
evaluation, primarily of insect remains. The procedures outlined in
57
Figure 4.1 Location of 7-15 Spurriergate site northeast of the River Ouse [red icon]. York Minister is shown at the
top of the photo and Clifford’s Tower is present in the bottom right corner (Google Earth 2009).
Kenward and associates (1980; 1985) were followed for the recovery of insect fossils from the
samples using paraffin floatation. The insect remains in the resulting residue and washover were
sorted using low-power binocular microscopes with the assistance of M. Rousseau, and identification
was performed through comparison with modern reference material in the collection of the former
Environmental Archaeology Unit, University of York.
The identifications were cross-checked
through the kindly assistance of H. Kenward.
4.2.3 Results and Analysis of the Data
The species identifications have been listed in Table 4.1, giving the minimum number of
individuals [MNI] represented by remains of each species for each sample. The nomenclature follows
Kloet and Hinck’s (1964; 1976; 1977; 1978) revised checklists for British Insects. A brief description
of each species’ modern habitat and diet is given along with the identifications.
Two taxonomic orders of insects, Diptera and Coleoptera, were recovered from the samples at
Spurriergate. The dipteran remains were primarily represented by pupae and were not identified.
However, the coleopteran remains were identified and for the most part taken to the generic if not
specific level. The Coleoptera were assigned to broad species groups based on ecology. In addition to
the species ecological groups outlined by Robinson (1981; 1983), a twelfth group was utilised, species
especially associated with stored grains.
This group is comprised of strong synanthropes, e.g.
Sitophilus granarius, which are associated with cereals and cereal products. The beetles are not
58
Table 4.1Species list for 7-15 Spurriergate, York
2000.584 6063 MNI per
Sample
Ecology
Taxon
4 6 7
Carabidae indet.
1
10 12
16
N/A
Dyschirius
(Hbst.)
1
on sandy loamy, clayey and muddy banks,
swamps, bogs, brick pits, moist arable fields,
loamy gardens, in leaves, moss, detritus,
flood debris, rotting vegetation, compost
(Koch 1989a).
globosus
1
Trechus
obtusus/quadristriatus
(Er./Schr.)
T. obtusus: in moist deciduous woodland
and on river banks, swamps, moist shaded
meadows. High mountains raised bogs.
Littoral wash zone. Under leaves and moss,
in detritus and Sphagnum sp. T.
quadristriatus: frequently synanthropic,
loamy arable fields, moist weedy areas,
gardens, woodland margins, hedges, shaded
banks, dunes, caves. In hay stacks and barns,
under rotting vegetation, in detritus and
compost (Koch 1989a)
1
1
prefers thick vegetation; loamy arable fields,
flood plains, meadows, woodland margins;
hedges and gardens; brick pits; gravel pits.
Littoral - wash zones. In decaying vegetation
and flood debris; under loose bark (Koch
1989a)
Pterostichus
melanarius (Ill.)
Hydraena
(Curtis)
2
in muddy streams or stagnant drainage
ditches (Duff 1993)
testacea
1
59
2000.584 6063 MNI per
Sample
Ecology
Taxon
4 6 7
10 12
16
H. aquaticus: in temporary pools and muddy
or weedy margins of ponds and lakes in
spring and autumn (Brown 1940). H.
grandis: in stagnant freshwater, preferring
eutrophic, more or less open and often
temporary pools with clayey and grassy
bottom (Hansen 1987)
Helophorus
aquaticus/grandis
(L./Ill.)
1
Helophorus sp.
1
Aquatic
Cercyon
haemorrhoidalis (F.)
Cercyon unipunctatus
(L.)
1
very eurytopic, in all kinds of decaying
organic matter, mainly in cow, horse and
sheep dung, but also frequently in rotting
plant debris, especially compost heaps, also
old mushrooms, flood debris on wetter
habitats, carrion, at sap, e.g. on birch, in
nests of various birds (Hansen 1987)
1 1 1
1
in all kinds of decaying organic matter,
distinctly synanthropic, in various debris
around farm buildings, e.g. compost heaps
and barn manure (Hansen 1987)
1
1
Cercyon atricapillus
(Marsham)
4 1 2
Cercyon terminatus
(Marsham)
1
3
in fields, weedy places, cow pastures, river
meadows, heaths, gardens and stables,
especially in fermenting materials (compost,
root heaps, stable manure, heaps of rotting
vegetation), fresh dung and carrion (Koch
1989a)
4
1
1
in herbivore dung or decaying grass (Duff
1993)
60
2000.584 6063 MNI per
Sample
Ecology
Taxon
4 6 7
10
12
16
associated with muddy detritus and decaying
plant litter in well vegetated stagnant waters,
including fens, marshes and swampy
overgrown canals, borrow pits and ditches
(Merritt 2006)
Cercyon tristis (Ill.)
Cercyon
(Payk.)
2 3
Cryptopleurum
minutum (F.)
Omalium
(Grav.)
in almost all rotting vegetation, in flood
debris and litter (Koch 1971)
analis
Megasternum
obscurum (Marsham)
Acritus
(Hoff.)
1
3
1
1
in decaying grass and herbivore dung (Duff
1993)
1 1 1
1
2
in all kinds of decaying organic matter,
usually very abundant, mainly in compost
heaps, rotting grass, in dung and at carrion,
also among various plant debris near water
(Hansen 1987)
1
1
in fields, meadows, gardens, weedy places,
woodland margins and pine heaths;
especially in the lower layers of old stable
dung heaps, in dung, rotting vegetation,
compost and rootcrop heaps, tannery waste,
wood debris and barn straw (Koch 1989a)
nigricornis
1
in grass tussocks, including cereal crops, or
in haystack refuse and leaf litter (Duff 1993)
caesum
Xylodromus
concinnus (Marsham)
1
in fields, river meadows, woodland margins
and woods. In straw in barns and stalls in
hay and compost heaps; occasionally also in
woods in wood mould and in nests in hollow
trees (Koch 1989a)
1 1
1
1
61
2000.584 6063 MNI per
Sample
Ecology
Taxon
4 6 7
Carpelimus bilineatus
(Steph.)
2 1 3
Oxytelus
(Grav.)
10
12
16
on sandy-muddy banks, fields, gardens,
woodland margins. Littoral: wash zone. On
mud, under detritus and flood debris, in wet
compost, straw and stable manure heaps
(Koch 1989a)
3
2
2
in fields; cattle pastures; weedy places;
gardens; woodland margins; stables. Littoral
- dunes.
In dung stable manure; in debris of
Phragmites sp.; in rotting hay (Koch 1989a)
sculptus
2 1 2
3
1
in rotting vegetation, also in fungi, in
mouldering leaves, litter and flood debris, in
game bird food debris, in compost, straw and
stable manure, in dung of cattle and man, in
bird nests and underground animal burrows,
on mud and in Sphagnum sp. (Koch 1989a)
Anotylus rugosus (F.)
1
1
in more or less fresh dung; in carrion; in
stable manure and compost heaps; in rotting
vegetation (Koch 1989a)
Anotylus sculpturatus
(Grav.)
Anotylus
(Grav.)
1
1
1
on damp soil, banks, swampy and wet
meadows, river meadows, copses and
gardens. Alpine - in pastures and green alder
zones. In rotting vegetation, also in fungi, in
stable manure and compost heaps (Koch
1989a)
nitidulus
1 1 3
2
1
In rotting vegetation, also fungi; in dung and
carrion; in compost and stable manure heaps;
in rotting straw in barns and ricks (Koch
1989a)
Anotylus
tetracarinatus (Block) 1
1
1
62
2000.584 6063 MNI per
Sample
Ecology
Taxon
4 6 7
Platystethus
arenarius (Geoff.)
2
Platystethus degener
(Muls. and Rey)
3
in herbivore dung and damp decaying
vegetation (Duff 1993)
1
1
in mud and damp litter near water (Duff
1993)
1
in mud and damp litter near water (Duff
1993)
1
in mud and damp litter near water (Duff
1993)
nitens
1
in meadows, woodland margins, woods,
fields and gardens. In rotting vegetation, in
straw debris, in straw and chaff in field barns
and heaps (Koch 1989a)
Lithocharis ochracea
(Grav.)
1
1
in woods, woodland margins, heaths,
especially in nests of Formica sp (Koch
1989a); in some numbers in haystack bottom
(Donisthorpe 1939)
Leptacinus
intermedius (Donis.)
Leptacinus
(Steph.)
Kateretes sp.
1
on field margins; weedy places; gardens;
woodland edges. Especially in old stable
manure heaps, in compost and rotting
vegetation; in rotting root crops; in rotting
marginal straw in barns and heaps (Koch
1989a)
pusillus
1
Philonthus sp.
Tachinus
(Grav.)
16
3
Platystethus cornutus
(Grav.)
Platystethus
(Sahl.)
10 12
1
1
1
1
1
N/A
in rotting vegetation, in compost and stable
manure heaps, in mouldy straw from barns
and heaps, in carrion and dung (Koch
1989a)
laticollis
1 1
4 3 4
5
2
3
N/A
63
2000.584 6063 MNI per
Sample
Ecology
Taxon
Monotoma
(Hbst.)
4 6 7
10 12
16
on field and meadow edges; gardens; weedy
places; rubbish dumps; In rotting and
mouldy vegetation, compost, stable manure,
hay, straw; occasionally on fungi, under dry
cattle dung and with ants (Koch 1989a)
picipes
1
in gardens; rubbish dumps; barns and stalls;
field and meadow edges; also woodland
margins and river meadows (Koch 1989a)
Monotoma longicollis
(Gyll.)
1 1
1
Oryzaephilus
surinamensis (L.)
18 10
Cryptolestes
ferrugineus (Steph.)
Cryptophagus
scutellatus (Newman)
2
in cereals and cereal products (Koch 1989a)
8 5 6
6
in cereals and their products; also in dried
fruit and peanuts; more rarely under more or
less dry bark of decaying and fallen trunks of
Fagus, Carpinus, Quercus, and Salix
viminalis, but also on conifers; occasionally
in mouldy straw in heaps and in leaf litter at
the foot of trees. Feeds on debris and mould
fungi;
also in deciduous and mixed
woodland; parks; river meadows; woodland
margins; field margins (Koch 1989b)
9 8 9
17 19
9
in stables, barns and cellars, gardens, field
and meadow edges, more rarely in stream
and river meadows (Koch 1989b)
1 1 4
2
1
64
2000.584 6063 MNI per
Sample
Ecology
Taxon
4 6 7
16
in rotting hay and straw, as well as in stable
manure and compost heaps, also in rotting
vegetation, beet heaps, fresh grass cuttings,
game food debris, dry dung and wild animal
droppings, more rarely in woodland litter,
detritus, grass tussocks and flood debris, as
well as at sap flows on trees (Koch 1989b)
Ephistemus globulus
(Payk.)
Lathridius
(grp.) (L.)
10 12
1
1
mycetophagous, in mould fungi; often
synanthropic, in barns, cellars, etc, in houses
on damp walls, in granaries & warehouses,
in rotting provisions, corn etc. In the wild,
among mouldy leaves, vegetation, near fungi
and tree fungi, Polyporus sp. (Horion 1961)
minutus
9 8 10 8
3
5
Enicmus sp.
1
1
2
1
4
Corticaria sp.
1 1 1
1
1
4
Mycetophagous
Mycetophagous
Typhaea
(L.)
Palorus
(Wiss.)
mostly synanthropic, in cellars, stalls, barns,
etc. in mouldy rotting materials, wood, hay,
straw, leaves, etc., often in provision stores
on corn etc in mills etc. Not a pest but a
mould feeder. In the wild, less common, on
tree fungi, in decaying fruit trees, amid
rotting leaves, etc. (Horion 1961)
stercorea
1 1
2
1
2
in mills and bakeries, also in deciduous
woods and woodland margins. Above all in
cereals, meal and bran supplies; also noted
under rotting bark of old deciduous trees,
particularly Fagus (Koch 1989b)
ratzeburgi
4 1 2
7
4
1
65
2000.584 6063 MNI per
Sample
Ecology
Taxon
Tenebrio
(F.)
4 6 7
10 12
in cellars, stalls, corn stores and mills, more
rarely in deciduous woodland, parks and
gardens. Above all in cereals and their
products, also in pigeon lofts and nests of
Passer domesticus, occasionally in hollow
deciduous trees, mostly in association with
old bird nests, under rotting bark, in mouldy
stumps (Koch 1989b)
obscurus
1
in sandy pastures, river floodplains and
woods, especially in fresh cattle and horse
dung, also in human faeces (Koch 1989b)
Aphodius
contaminatus (Hbst.)
Aphodius
(L.)
Aphodius
(L.)
16
1
in all dung, more rarely in compost and
stable manure heaps and rotting vegetation,
especially cabbage stalks, and human faeces
(Koch 1989b); known to be a pest of
potatoes and cultivated mushrooms (Jessop
1986)
fimetarius
1
in cattle pastures; fields; stream and river
meadows; vineyards and gardens; woodland
margins; heaths. in rotting and fermenting
vegetation, rotting beets, cabbage stalks,
grape husks, also in silage heaps, compost
and stable manure heaps, as well as in dung
of pets and humans; occasionally on carrion
(Koch 1989b)
granarius
Phyllopertha
horticola (L.)
2 1
2
1
on meadows, woodland margins, hedges and
gardens, river floodplains, fields and weedy
places, on deciduous trees and shrubs, also
on fruit trees; larvae on roots of grasses, also
cereals, and clover (Koch 1989b)
1
66
2000.584 6063 MNI per
Sample
Ecology
Taxon
Hoplia
(Fues.)
4 6 7
10 12
16
on sandy river floodplains and banks of
rivers and lakes, heaths, part dry lawns,
swarms in the morning; especially on small
leaved Salix spp., but also on fruit trees, on
twig ends of young Pinus, on cereals and
flowers, Umbelliferae and Spiraea (Koch
1989b)
philanthus
1
1
in damp meadows; stream and river
meadows; field and meadow margins; arable
fields; woodland margins and wood pasture
(Koch 1992); larvae and adults on Rumex sp.
especially R. obtusifolius and R. crispus
(Bentley and Whittaker 1979; Chuter 2000)
Gastrophysa viridula
(Deg.)
1
in swamps and bogs, swampy banks and
meadows, copses (Koch 1992)
Prasocuris
phellandrii (L.)
1
N/A
Galerucella sp.
1
on Cruciferae, often on disturbed or
cultivated ground and a pest of cultivated
turnip, Brassica sp. (Duff 1993)
Phyllotreta nemorum
(L.)
Chaetocnema
concinna (Marsham)
2
oligophagous on Polygonaceae, more rarely
on Beta, Fagopyrum and Rheum; in winter,
singly in leaves, twigs, moss, detritus, hay,
straw, rotting vegetation and flood debris;
larvae in the shoot leaf base region (Koch
1992)
1
in fields, often Leguminosae fields, weedy
places, causeways and embankments, stream
and river meadows, dry and part dry lawns,
gardens and city parks, more rarely swampy
meadows
Sitona lineatus (L.)
1
67
2000.584 6063 MNI per
Sample
Ecology
Taxon
4 6 7
10 12
16
and bogs. Oligophagous on very many
Fabaceae, rarely also on Robinia sp., in
winter singly in leaves, hay, straw, grass
tussocks, litter, rotting vegetation, moss on
trunks, also in flood debris (Koch 1992); a
serious pest of peas (Jones and Jones 1974)
on heaths and bogs; dry woodland margins,
oligophagous on Betula sp. (Koch 1992)
Magdalis carbonaria
(L.)
Sitophilus
(L.)
Mecinus
(Hbst.)
1
lives and develops in grains of corn, rye,
barley, maize, oats, buckwheat, millet,
chickpeas, more rarely in chestnuts, acorns
corn meal (Hoffmann 1954)
granarius
3 1 1
5
2
on dry slopes, heaths, dry field margins,
sunny stream and river meadows, sunexposed woodland margins. Oligophagous
on Plantago spp., especially P. lanceolata;
occasionally in moss (Koch 1992)
pyraster
Gymnetron
pascuorum (Gyll.)
4
1
in degraded marshy grassland (Anderson
1998); on dry and warm slopes; dry swards;
dry meadow edges. Monophagous on
Plantago lanceolata ( Koch 1992)
1
References for species’ habitats and diets cited in Buckland and Buckland 2006
known to infest cereals in the field and a typically considered to indicate stored products. Moreover,
the 7-15 Spurriergate samples did not contain any species associated with Species Group 10, species
especially associated with structural timbers.
Approximately 95 % of the coleopteran remains have been classified into one of the ecological
categories. The species groupings are presented in Figures 4.2- 4.3. Figure 4.2 shows the number of
species recorded for each ecological group as a percentage of the assemblage. In Figure 4.3, the
minimum number of individuals per ecological is displayed.
68
Figure 4.2 Number of species by ecological group
100%
Stored grain
Grass roots
Lathriidiidae
Syanthropic
% Number of Species
80%
60%
Foul Organic
Wood/ trees
Meadowland
Disturbed ground/ arable
40%
Marsh/ aquatic plants
Dung/ Pastureland
Aquatics
20%
0%
Figure 4.3 Minimum number of individuals by species group
100%
% Number of Individuals
90%
Stored grain
Grassroots
80%
70%
Lathridiidae
Synanthropic
Foul organic
Wood/ tree
60%
50%
Meadowland
Disturbed ground/ arable
Marsh/ aquatic plant
40%
30%
Dung/ Pastureland
Aquatic
20%
10%
0%
4.2.4 Palaeoenvironmental Reconstruction of Roman Spurriergate
The aquatic and waterside environment
Water and marsh-related beetles comprised approximately 17 % of the coleopteran species
from Spurriergate, York with the aquatic insect fauna only forming around 3.4 % of the entire
assemblage.
Although there was diversity in the species reported, the number of individuals
constituted only 10 % of the assemblage. The aquatic beetles were largely associated with stagnant or
69
muddy water. Helophorus aquaticus/grandis species, in particular, are common in stagnant pools of
water. The species Hydraena testacea and Cercyon tristis are found in detritus on the muddy margins
of well-vegetated ponds and sluggish drains (Friday 1988; Merritt 2006), and the beetles provide
evidence towards the rich waterside fauna from the samples. Several Hydraenidae species readily
leave water and have been recorded on the muddy banks of the watersides. The Spurriergate samples
revealed several non-aquatic species that frequent muddy environments. The staphylinids Platystethus
degener, P. cornutus, and P. nitens are species that are found in environments comprised of highly
organic mud and decaying litter such as near ponds and rivers (Hammond 1971; Duff 1993). Flood
zone species of Carabidae were also noted in the assemblage. Pterostichus melanarius is a predaceous
ground beetle that hunts in decaying vegetation and debris in wash zones (Koch 1989a).
The water and marsh- associated fauna paint a picture of a potentially fetid environment with
stagnant pools of water.
There appears to have been rich layer of vegetation, though perhaps
decaying, overlaying or intermixed with a muddy or damp-disturbed matrix. Despite the site’s
proximity to the River Ouse (approximately 83 metres), there was no evidence of species associated
with flowing water and little indication of species which are phytophagous on waterside vegetation.
However, Gastrophysa viridula was present in small numbers and has been noted living on Rumex sp.
near marshes, and Prasocuris phellandrii seems to be oligophagous on aquatic Umbelliferae such as:
Oenanthe phellandrium, O. aquatica, O. crocata, Cicuta virosa, and Sium latifolium (Koch 1992;
Bullock 1993). Although phytophagous taxa were scarce, the site did contain a number of species that
would likely have been autochthonous to the muddy environments near the riverbanks.
The woodland and scrub
Wood and tree-related species made up less than 1 % of the Coleoptera from the first century
AD Spurriergate contexts. This estimate is comprised of beetles associated with Group 4 of the
species’ ecological groupings discussed above. While other species in the assemblage may contribute
to a woodland community, this category represents fauna associated with the bark, wood, leaves, etc.
of trees and shrubs rather than other forest elements such as litter or woodland herbs. The Group 4
fauna make up a minute portion of the assemblage. However, if Cryptolestes ferrugineus and Palorus
ratzeburgi, which are considered primarily to be grain pests but have been recorded under tree bark,
are added to Group 4, the number of individuals that are capable of being inhabiting trees increases to
23 % of the assemblage. The two of the species are fairly poor indicators of the presence of trees, and
their abundance in the samples is more likely associated with stored cereals [see Chapter 5]. The
scarabaeoid Hoplia philanthus has been recorded on Salix sp. (willow) and Pinus sp. (pine), especially
on river banks, meadows, and heaths (Koch 1989a). However, the species has also been noted on
cereals and flowers, and its larvae feed on grass roots. The curculionid Magdalis carbonaria is
perhaps the only true representative of a Group 4 species in the samples. The weevil is fairly host70
specific and may be a strong indicator for the presence of Betula sp. (birch). The larvae of M.
carbonaria feed on the interior of dead branches and twigs of birch (Hyman 1992). As Magdalis
carbonaria is only represented by a single specimen, MNI =1, from Sample 7, the presence of trees
near or on the site is not strongly supported by the insect remains.
Grassland, arable, and the open environments
The Early Roman landscape around the Spurriergate site, or perhaps more accurately around
York, was largely open. The region seemed to support grassland and/or pastureland. 18 % of the
archaeologically recovered terrestrial Coleoptera and 4 % of the individuals fall in the Group 3
category, indicating the presence of meadowland. Moreover, there were a few species whose larvae
feed on grassroots. The chafer Phyllopertha horticola as well as the Hoplia philanthus and other root
feeding scarabaeoids were present in the samples but not abundant. The phytophagous fauna of
grassland species is representative of the presence of trefoils and flowery herbs.
A few of the more host-specific phytophagous beetles from the samples and their favoured
plant foods include:
Gymnetron pascuorum
Plantago lanceolata
Mecinus pyraster
P. lanceolata and P. media.
While considered a pest of peas, the pea-leaf weevil Sitona lineatus has been recorded on
Fabaceae, Lathyrus spp., Pisum sativum, Pisum spp., Trifolium spp., Vicia faba, and Vicia spp.
(Bullock 1993; Morris 1997).
Moreover, its larvae are oligophagous root feeders.
The single
specimen of S. lineatus from Spurriergate may hint at the presence of meadowland or agricultural land
in the vicinity of the site.
Along with the pea-leaf weevil, 5 % of the Coleoptera species from the samples support the
presence of disturbed or arable land. There were a few individuals of Phyllotreta nemorum, which is
associated with various Cruciferae such as the turnip Brassica sp. An individual of Chaetocnema
concinna was also recovered. While C. concinna feeds on various cultivated Polygonum sp. and
Rumex sp. (see LeSage 1990 for a comprehensive list), it could also represent grassland or waterside
environments (Richards 1926; Duff 1993). While primarily coprophilous, Aphodius fimetarius is also
considered a pest of potatoes and cultivated mushrooms (Jessop 1986).
Several of the coleopteran species are representative of hay or straw. For example, Leclercq
(1946) found numerous Typhaea stercorea in hay brought in from the meadow. However, the beetle
is more commonly noted in mouldy or decaying hay refuse. Several other phytodetriticolous species,
which are loosely associated with decaying hay and dry vegetable matter, from Groups 7 and 9, were
recovered from the Spurriergate samples. Ephistemus globulus, Trechus quadristriatus, Omalium
caesum, and Xylodromus concinnus were prevalent in the samples. An individual specimen of Acritus
nigricornis which is common in ‘sweet’ compost such as hay, straw, and cut grasses, was also found.
71
The site also contained several other decomposer species, which while not particularly associated with
hay does indicate the presence of vegetable refuse.
Whereas these species are associated with
vegetation, their association with decaying matter probably is a reflection of conditions on the site
rather than the surrounding environment.
Along with the vegetation decomposer fauna, Group 7 was comprised of dung indicators. The
foul organic-associated species constitute 21.5 % of the assemblage’s species and 14.5 % of the
individual specimens. However, most of Group 7 beetles will feed on decaying vegetation as well as
herbivore dung, which is essentially another form of decaying vegetation. A better indication of the
presence of dung is Species Group 2. 7 % of the recovered Coleoptera specimens are associated with
dung and pastures. In general, Group 2 dung beetles are fairly strong indicators of the presence of
dung as the species tend to burrow in or under patties. The recovery of Group 2 species at an MNI of
5 % suggests the presence of the herbivorous mammals at the site.
The dung-feeding beetles are not typically associated with the excrement of a single species.
Megasternum obscurum was present in amble numbers and provides a more general indication of the
presence of herbivore dung. Aphodius granarius and Cercyon haemorrhoidalis are common in cow
patties but have also been found in the manure of other domesticated herbivores such as sheep and
horses. Cercyon atricapillus is mainly associated with horse and cattle excrement. Hansen (1987)
proposed that Cercyon terminatus was mainly found in cattle and horse dung whereas Koch (1989a)
put forth that the species was particularly associated with sheep and cattle manure. Most of the dung
fauna are associated with domesticated species; however, Donisthorpe (1939) suggests that Aphodius
contaminatus is more common in deer excrement. A. contaminatus was only evidenced by a single
specimen at the site, and the beetle has also been recorded on old horse and cattle dung (Landin 1961).
The Spurriergate samples were comprised of a fair proportion of meadowland, arable, and foul
organic species. Although the Coleoptera may suggest that agricultural land and grassland used for
grazing were in the vicinity of first century AD Roman York, the MNI for the groups indicates that the
fauna, while present, were not abundant. In comparison, the beetles, which were associated with
decomposing vegetation and dung, were strongly represented. In an open environment, one would
expect the presence of phytophagous and root-feeding fauna to be more pronounced in comparison to
the rotting vegetation associates. The abundance of dung beetles in association with the synanthropic
hay species such as Typhaea stercorea suggest the existence of a more confined environment that is
suitable for maintaining domesticated herbivore mammals, i.e. a stable or barn, on the site.
Other habitats
There were several species associated with human habitation in the Spurriergate contexts. In
addition to the synanthropic Coleoptera assigned to Group 9 (discussed above), 39 % of the
individuals that were recovered from the samples were species which are strongly associated with the
72
presence of stored cereals. Cryptolestes ferrugineus and Oryzaephilus surinamensis comprise part of
the grain fauna and were abundant in all the samples.
obscurus was also present.
Additionally, an individual of Tenebrio
The species Sitophilus granarius has only been recorded from
synanthropic environments in temperate regions; the granary weevil is strongly associated with stored
cereals and does not appear to infest grains that are still in the field [see Chapter 5]. The recovery of
significant numbers of S. granarius from the site is a good indicator of the presence of cereal grains.
While Palorus ratzeburgi is also an indicator species for stored cereals, the species may also provide
evidence of the condition of the grains. The small-eyed flour beetle is common on mouldy or
damaged grains (Brendell 1975).
Group 8 associates were also prevalent in the samples. Lathridius minutus group, in particular,
was abundant and constituted 10.5 % of the individuals from the Spurriergate assemblage. Lathridius
minutus group species are typical but non-obligate synanthropes. The beetles are mycetophagous and
are believed to feed on mould, spores, and hyphae (Larsson and Gigja 1959). Although the species
have been noted in plant debris in birch forests (Böcher 1988) and in Rumex sp. in meadows
(Bengtson 1981), L. minutus group is more frequently associated with hay barns, stables, and granaries
(Horion 1961; Lindroth et al. 1973; Barker and Smith 1990).
4.2.5 Palaeoclimatic Reconstruction
Sample
TMaxLo
TMaxHi
TMinLo
TMinHi
TRange
Lo
TRange
Hi
NSPECIES
Overlap
Table 4.2 Mutual Climatic Range method predictions for Roman 7-15 Spurriergate, York
4
6
7
10
12
16
Site
11
10
15
15
10
15
15
18
11
18
25
18
26
18
-17
-6
-14
-11
-12
-13
-9
6
-3
7
10
7
11
6
11
14
11
9
11
9
11
30
16
29
28
24
32
24
7
6
6
6
5
4
15
100
100
83.33
83.33
80
100
93.33
(calculated using Buckland and Buckland 2006)
Table 4.2 and Figure 4.4 show the climate predictions calculated using the BugsMCR program
(Buckland and Buckland 2006). Only carnivorous and scavenging species were analysed, as the
ranges of herbivorous or phytophagous Coleoptera may only reflect the distribution of their food
plants rather than provide a ‘true’ climatic signal. Fifteen species were evaluated [Appendix 1A], and
predicted the temperature of the warmest month as ranging between 15 ˚C and 18 ˚C and the
temperature for the coldest month between -9 ˚C and 6 ˚C. This range of temperatures is similar to the
73
estimates evidenced by the beetles from other Romano-British sites such as Bedern Well, York,
Alcester, Warwickshire, and Copthall Avenue, London [Appendix 1B].
Figure 4.4 7-15 Spurriergate, York MCR estimates by sample
30
25
TMax
20
15
10
5
2000.584.6063_16
2000.584.6063_12
2000.584.6063_10
2000.584.6063_7
2000.584.6063_6
15
10
5
0
-5
-10
-15
-20
2000.584.6063_4
TMin
0
(calculated using Buckland and Buckland 2006)
4.2.6 Discussion: The Environment and Climate of 7-15 Spurriergate and Implications for
Culture Contact
The Early Roman insect assemblages from Spurriergate are interesting because they consist of
strongly synathropic species as well as grassland and riverside fauna. Kenward and Hall (1997) have
proposed that the presence of grain pests along with ‘hay’ fauna, house fauna, and decomposers is
characteristic of stable manure. Although the Spurriergate samples evaluated here did not yield the
characteristic spider beetles, Ptinus fur and Tipnus unicolor, which comprised part of Kenward and
Hall’s (1997) stable manure indicator group, the ecological association evidenced by the assemblage
may infer a stable fauna.
The grains may have served directly as a part of the mammals’ diet or, less possibly, the grain
pests could have invaded residue grain in straw or chaff that was used for bedding. Moreover,
Osborne (1983) demonstrated that insect remains can successfully pass through a human dietary tract
without damage; it seems plausible that the same would hold true for large non-ruminant herbivores.
The presence of meadowland fauna could have been ingested during grassland grazing and introduced
to the urban site through the dung of the herbivores or may have been imported alongside vegetation
74
used for floor litter. The site produced a range of fauna associated with plant debris, in various stages
of decay. Furthermore, the samples yielded numerous dung beetles associated with large herbivores as
well as coleopteran species reflecting the presence of hay. If viewed independently, the species in the
assemblage could represent a number of habitats; however, collectively they strongly infer a stable or
barn environment. The aquatic and waterside fauna, which perhaps do not categorise with the stable
manure indicator group, were present in low numbers, and may stand as a background fauna attracted
to the muddy micro-habitat from the nearby River Ouse.
While most of the species are representative of a stable or barn environment, a few of the
Coleoptera may also reflect local or foreign exchange.
The grain fauna and the hay fauna, in
particular, may provide evidence towards palaeoeconomic activities in Roman York as the species
may reflect commodities which are transported and/or traded by man. Using of the palaeoecological
approach, the potentially heterochthonous species will be identified by comparing the species’
temperature requirements for completion of life cycle to the estimated MCR for the site at TMax 18
˚C, which is three degrees lower than the average TMax for the warmest month in modern York (cf.
WatkinsHire n.d.).
The grain fauna was comprised of species requiring a range of temperatures in order to carry
out their developmental life cycle [see Figure 5.2 for a detailed comparison]. Based on temperature
requirements, Tenebrio obscurus (14-30 ˚C) and Sitophilus granarius (15-35 ˚C) slightly overlap with
the TMax of the assemblage, which implies that the species may have been capable of surviving and
completing their development during the warmer months outside of human habitation in Roman
Britain. However, both species have optimal temperature ranges that are significantly higher than the
calculated maximum temperature for the warmest month. This suggests that the species are not
indigenous to Britain and would have most likely been introduced at some point. The minimum
temperature requirements for the other grain-associated species equal or exceed 18 ˚C. As such, the
pests would not have been able to complete their development in the wild and are unlikely to have
been native. However, the species would have likely been able to reproduce and develop in Roman
Britain under synanthropic conditions. Granaries and fermenting vegetation would have had higher
temperatures than their surrounding environment, which would have enabled the species, once
introduced, to become established.
Solely on the basis of temperature requirements and
palaeoclimatic reconstructions, the grain fauna, as a whole, suggest the importation of cereals from
warmer regions by the Romans.
The hairy fungus beetle Typhaea stercorea is strongly associated with hay. While the species
does not appear to damage the hay directly, as it is mycetophagous, it seems associated with the fungi
and mould common to sweet compost. Laboratory assessments by Jacob (1988) have demonstrated
that T. stercorea is capable of completing its development at 17.5 to 30 ˚C when held at 90 % relative
humidity. At 15 ˚C, the eggs did not hatch and the pupa did not develop. Similar to the majority of
75
the grain fauna, the hairy fungus beetle may have been able to endure and become established in
synanthropic environments but would not have been capable of surviving in the wild during the first
century of the Roman occupation of Britain.
The presence of the species implies an original
connection to a warmer climate.
4.2.7 Summary
The samples evaluated from 7-15 Spurriergate York yielded Coleoptera associated with a
range of habitats. When the ecologies were viewed collectively, the fauna appeared to reflect a stable
or barn environment used to keep herbivorous mammals. The presence of meadowland and cereal
beetles may indicate that the domesticated animals had a diet that involved pastureland grazing and
grain supplements. While the majority of the species were likely associated with the local and
hinterland environments around Roman York, the temperature ranges for the grain beetles and
Typhaea stercorea indicate warmer climate origins for those species and suggest the potential
introduction of those Coleoptera to the site, which would infer the importation of cereals and hay to
York. However, the species would have most likely been able to survive in Britain under the
increased temperatures presented by their synanthropic environments. As the present palaeoecological
study does not consider other archaeoentomological evidence which is necessary to assess the spatial
or temporal changes in species’ distributions (that being considered with the biogeographical approach
in Chapter 5), it is not possible to discern how early the grain beetles and hay associates arrived in
York.
4.3 Case Study: 16-22 Coppergate, York Period 4b
4.3.1 Introduction
The archaeoentomological assessments regarding 16-22 Coppergate, York were carried out as
part of a series of large-scale urban rescue excavations in York, which were conducted by the York
Archaeological Trust during the 1970s and 1980s. The 16-22 Coppergate site is situated within York
city centre approximately 200 m from the River Foss and 100 m from the River Ouse [Figure 4.4].
The site yielded contexts ranging in date from the mid-9th (cf. Hall and Kenward 1999a) to the early
11th century (cf. Hall and Kenward 1999c).
4.3.2 Background and Processing Methods
The insect remains were processed and identified following the procedures outlined in
Kenward (1985; 1992). An account of the non-vertebrate biological remains, which were recovered
76
from the site, has been published by Kenward and Hall (1995). During the original processing and
identification period, the
Figure 4.4 Location of 16-22 Coppergate site (near red icon). The site falls between the River Foss (right) and the
River Ouse (left) near the Jorvik Viking Center. Clifford’s Tower can be seen at the bottom middle of the photo
(Goggle Earth 2009).
Figure 4.5 Post-and-wattle structures
(Richards 1991, 67)
77
investigators employed a range of recording methods: detail recording, scan recording, semiquantitative scan recording, and non-qualitative rapid scan recording (Kenward and Hall 1995). In
this study, insect fossils from five samples were selected for further evaluation. The materials relate to
contexts 22574 and 22490, which were dated between 1020 to 975 BP (Period 4b; cf. Hall and
Kenward 1999b). The contexts are believed to be associated with the Anglo-Scandinavian Period.
The remains of four tenements were found at the site, dated to Period 4b. The tenements,
which are portrayed in Figures 4.5-4.7, were delineated by
Figure 4.6 Period 4B tenements
wattle fences comprised mainly ofoak and hazel and were
categorised as the single post-and-wattle style (O’Connor
1994). The five samples were retrieved from fills inside the
‘west wicker building’, Tenement C (Hall and Kenward
1999b). Context 22574 was comprised of very dark grey,
silty-sandy clay, which contained lenses of brownish-yellow
to olive-yellow compacted grass/straw and circa 10 % wood
chips. Context 22490 was excavated from the uppermost
fill; it was predominately a black, peaty, silty- clay loam,
comprised of approximately 15 % wood chips (Hall and
Kenward 1999b).
4.3.3 Results and Analysis of Data
Table 4.3 provides a detailed species list for contexts
22574 and 22490 and indicates the determined minimum
(Hall 1984, 50)
number of individuals [MNI] of each species for each
sample. Kloet and Hinck’s (1964; 1976; 1977; 1978) revised checklists for British Insects was
employed for species’ nomenclature. Table 4.3 also provides a brief description of each species’
modern habitat and diet.
Six taxonomic orders of insects were recovered from contexts 22754 and 22490: Coleoptera,
Diptera, Anoplura, Mallophaga, Siphonaptera (fleas), and Hymenoptera. The insect remains were
identified to the generic if not specific level. The ecological coding system proposed by Robinson
Figure 4.7 Drawing of post-and-wattle houses
(Murray 1983, vi)
(1981; 1983) was utilised to assign the Coleoptera to broad species groups. An additional ecological
group was added to denote species especially associated with carcasses. The species associated with
the remaining orders were grouped as either ectoparasites or bees.
77 % of the coleopteran species have been categorised into one of the ecological categories.
The species groupings are presented in Figures 4.8- 4.10. Figure 4.8 shows the number of species
recorded for each ecological group as a percentage of the whole assemblage. Figure 4.9 expresses the
results as the minimum number of individuals per species group, and Figure 4.10 displays the MNI
values of the coleopteran fauna as a percentage of the beetle assemblage. Figure 4.11 does not include
non-coleopterous species for clarification purposes as the hymenopteran individuals alone comprised
74.5 % of the whole assemblage, which rendered the other groups largely indiscernible in the chart.
With the non-coleopterous individuals included, the MNI of only four of the species groups, in
addition to the Bees, constituted greater than 1 % of the entire assemblage: Meadowland 1.1 %,
Dung/foul organic material 5.1 %, Lathridiidae 2.7 %, and Synanthropes 3.1 %.
79
Table 4.3 Species list for selected contexts from 16-22 Coppergate, York
Ecology
1469/T
22490
MNI
per
sample
1469/1
1486/T
1486/1
22574 MNI
per sample
1485/1
Taxon
Coleoptera
Carabidae
indet.
1
fossor
a eurytopic species, usually in
open country on rather humic
ground with more or less
dense vegetation of grasses;
feeds on both vegetable and
animal matter, including
larvae
and
pupae
of
Meligethes
sp.
(Lindroth
1985)
2
Clivina
(L.)
1
1
N/A
Trechus micros
(Clair.)
1
usually near water, but also in
damp grassland, probably
associated with runs of small
mammals (Luff 1998); syn.
Trechoblemus micros Hbst.
Trechus sp.
1
moist vegetation, usually near
water
Bembidion
gilvipes (Sturm)
1
in moist meadows, moist water
meadows, woods, swampy
woods, swampy banks. Under
detritus
and
rotting
vegetation, in flood debris, in
grass (Koch 1989a)
Bembidion
biguttatum (F.)
1
in eutrophic fens bordering
lakes and rivers, usually
among tall vegetation; moss
and leaves at margins of
ponds and pools (Lindroth
1985)
80
Ecology
1469/T
22490
MNI
per
sample
1469/1
1486/T
1486/1
22574 MNI
per sample
1485/1
Taxon
Harpalus
rubripes (Duft.)
1
dry arable fields with sparse
vegetation; dry meadows,
woodland
margins
and
clearings; sandy river banks
and coasts.
Under grass
tussocks, leaves and moss
(Koch 1989a)
Pterostichus
melanarius (Ill.)
1
prefers
thick
vegetation;
loamy arable fields, flood
plains, meadows, woodland
margins; hedges and gardens.
In decaying vegetation and
flood debris; under loose bark
(Koch 1989a)
Amara sp.
1
species in bare ground and
arable
on
sandy
soils
(Robinson 1991)
Hydropor-inae
indet.
1
Aquatic
Ochthebius sp.
1
3
Aquatic
Aquatic
1
Sphaeridium sp.
1
1
Helophorus sp.
open-grazed land; dung
Cercyon
haemorrhoidalis (F.)
1
2
3
very eurytopic, in all kinds of
decaying organic matter,
mainly in cow, horse and
sheep
dung,
but
also
frequently in rotting plant
debris, especially compost
81
Ecology
1469/T
22490
MNI
per
sample
1469/1
1486/1
1486/T
22574 MNI
per sample
1485/1
Taxon
heaps, also old mushrooms,
flood debris on wetter
habitats, carrion (Hansen
1987)
1
3
Cercyon
unipunctatus
(L.)
Cercyon
atricapillus
(Marsham)
in all kinds of decaying
organic matter, distinctly
synanthropic,
in
various
debris around farm buildings,
e.g. compost heaps and barn
manure (Hansen 1987)
1
in fields, weedy places, cow
pastures, river meadows,
heaths, gardens and stables,
especially
in
fermenting
materials
(compost,
root
heaps, stable manure, heaps of
rotting vegetation), fresh dung
and carrion (Koch 1989a)
Cercyon
terminatus
(Marsham)
1
in herbivore dung or decaying
grass (Duff 1993)
in
almost
all
rotting
vegetation, in flood debris and
litter (Koch 1971)
1
4
6
Cercyon analis
(Payk.)
Megasternum
obscurum
in decaying grass and
herbivore dung (Duff 1993)
1
1
(Marsham)
Cryptopl-eurum
minutum (F.)
1
in all kinds of decaying
organic matter, usually very
abundant, mainly in compost
heaps, rotting grass, in dung
and at carrion, also among
various plant debris near
water (Hansen 1987)
82
Ecology
1469/T
22490
MNI
per
sample
1469/1
1486/1
1486/T
22574 MNI
per sample
1485/1
Taxon
Hydrobius
fuscipes (L.)
1
very eurytopic, mainly in
stagnant water, but also in
slower reaches of streams,
both fresh and brackish water.
Usually among vegetation in
shallows near water's edge
(Hansen 1987)
Histeridae
indet.
1
dung and carrion
in fields, meadows, gardens;
especially in the lower layers
of old stable dung heaps, in
dung,
rotting
vegetation,
compost and rootcrop heaps,
tannery waste, wood debris
and barn straw (Koch 1989a)
1
1
Acritus
nigricornis
(Hoff.)
Orthoperus sp.
1
3
Moulds
Ptenidium sp.
1
1
rotting vegetation
N/A
2
3
Acrotrichis sp.
Dropephylla sp.
1
fungi under bark
Omalium
rivulare (Payk.)
1
in grass tussocks including
cereal crops, or in damp litter,
fungi or carrion (Duff 1993)
83
in fields, river meadows,
woodland margins and woods.
In straw in barns and stalls in
hay and compost heaps;
occasionally in wood mould
and in nests in hollow trees
(Koch 1989a)
1
9
Xylodromus
concinnus
(Marsham)
2
Ecology
1469/T
22490
MNI
per
sample
1469/1
1486/1
1486/T
22574 MNI
per sample
1485/1
Taxon
Omaliinae
indet.
1
N/A
on sandy-muddy banks, fields,
gardens, woodland margins.
Littoral: wash zone. On mud,
under detritus and flood
debris, in wet compost, straw
and stable manure heaps
(Koch 1989a)
1
1
1
Carpelimus
bilineatus
(Steph.)
Carpelimus
fuliginosus
(Grav.)
1
in fields; gardens; weedy
places. Subalpine pastures. In
compost, stable and rotting
hay heaps (Koch 1989a);
associated with wetlands, in
damp and marshy places
under leaves, in moss,
amongst herbage and in
tussocks (Hyman 1994)
Carpelimus
pusillus (Grav.)
1
in gardens, fields, weedy
places, also on sandy and
swampy banks. In stable
manure and compost heaps, in
dung
beds;
in
rotting
vegetation, in rotting straw
and field barns and ricks,
under grass tussocks and
debris (Koch 1989a)
84
Ecology
1469/T
22490
MNI
per
sample
1469/1
1486/1
1486/T
22574 MNI
per sample
1485/1
Taxon
Carpelimus sp.
1
fetid environments
1
2
Oxytelus
sculptus (Grav.)
1
1
Anotylus
rugosus (F.)
Anotylus
sculpturatus
(Grav.)
1
1
on damp soil, banks, swampy
and wet meadows, river
meadows.
In
rotting
vegetation, also in fungi, in
stable manure and compost
heaps (Koch 1989a)
in cut grass, fungi, damp
straw, and moss (Donisthorpe
1939)
1
6
2
1
in rotting vegetation, also in
fungi, in mouldering leaves,
litter and flood debris, in
compost, straw and stable
manure, in dung of cattle and
man, in bird nests and
underground animal burrows,
on mud and in Sphagnum sp.
(Koch 1989a)
in more or less fresh dung; in
carrion; in stable manure and
compost heaps; in rotting
vegetation (Koch 1989a)
Anotylus
nitidulus
(Grav.)
Anotylus
complanatus
(Er.)
in fields; cattle pastures;
weedy
places;
woodland
margins; stables. In dung
stable manure; in debris of
Phragmites sp.; in rotting hay
(Koch 1989a)
Anotylus
in rotting vegetation, also
fungi; in dung and carrion; in
compost and stable manure
heaps; in rotting straw in
barns and ricks (Koch 1989a)
tetracarinatus
1
(Block)
85
Ecology
Platystethus
arenarius
(Geoff.)
1
Platystethus
nitens (Sahl.)
1
1469/1
1469/T
22490
MNI
per
sample
in mud and damp litter near
water (Duff 1993)
2
1486/1
1486/T
22574 MNI
per sample
1485/1
Taxon
N/A
1
in herbivore dung and damp
decaying vegetation
(Duff
1993)
2
Stenus sp.
Lithocharis
ochracea
(Grav.)
1
in
meadows,
woodland
margins, woods, fields and
gardens. In rotting vegetation,
in straw debris, in straw and
chaff in field barns and heaps
(Koch 1989a)
Leptacinus
pusillus (Steph.)
2
on field margins; weedy
places; woodland edges. in
old stable manure heaps, in
compost
and
rotting
vegetation; in rotting root
crops; in rotting marginal
straw in barns and heaps
(Koch 1989a)
Gyrohypnus
fracticornis
(Müll.)
1
in rotting vegetation, also in
fungi, in compost and stable
manure heaps, in marginal
straw in barns and heaps, in
carrion, in detritus and flood
debris (Koch 1989a)
Gyrohypnus sp.
1
1
fetid environments
Xantholinus
linearis (Ol.)
1
in grass tufts, haystack refuse,
dead leaves, at roots of
heather (Buck 1955)
86
Ecology
1469/T
22490
MNI
per
sample
1469/1
1486/1
1486/T
22574 MNI
per sample
1485/1
Taxon
Philonthus sp.
3
1
1
N/A
Staphylininae
indet.
3
N/A
Tachyporus sp.
1
1
N/A
Tachinus rufipes
(L.)
1
in decaying grass and grass
tussocks or at plant roots,
including arable crops, and in
herbivore dung in woods and
grassland (Duff 1993)
in rotting vegetation, also
fungi, in compost, stable
manure, hay and straw heaps,
also in dung and carrion,
under stones and in grass
tussocks, in detritus (Koch
1989)
5
1
Cordalia
obscura (Grav.)
Crataraea
suturalis
(Mann.)
3
in vicinity of dwellings, in
mouldy straw in barns, sheep
pens and cellars, also in nests
(Harde 1984)
87
Ecology
1469/T
22490
MNI
per
sample
1469/1
1486/1
1486/T
22574 MNI
per sample
1485/1
Taxon
Aleocharinae
indet.
1
2
7
N/A
Pselaphidae
indet.
1
1
N/A
Euplectini indet.
1
N/A
Melanotus sp.
1
rotting wood and bark
Cyphon sp.
1
near water
Dermestes sp.
1
1
animal carcasses
Brachypterus sp.
1
on Urtica sp. (nettles)
88
Ecology
1469/T
22490
MNI
per
sample
1469/1
1486/1
1486/T
22574 MNI
per sample
1485/1
Taxon
Meligethes sp.
1
1
1
meadowland vegetation
Omosita sp.
1
2
carrion
Monotoma
spinicollis
(Aubé)
1
in rotting and mouldy
vegetation,
especially
compost, hay and straw; also
in old stable manure heaps
(Koch 1989a)
Monotoma
picipes (Hbst.)
1
on field and meadow edges;
gardens;
weedy
places;
rubbish dumps; in rotting and
mouldy vegetation, compost,
stable manure, hay, straw;
occasionally on fungi, under
dry cattle dung and with ants
(Koch 1989a)
Monotoma
bicolor (Villa)
3
in plant debris (Duff 1993);
rotting vegetation, compost,
hay (Koch 1989a)
Cryptophagus
1
1
scutellatus
(Newman)
in stables, barns and cellars,
gardens, field and meadow
edges, more rarely in stream
and river meadows (Koch
1989b)
Cryptophagus sp.
1
1
9
1
N/A
89
Ecology
1469/T
22490
MNI
per
sample
1469/1
1486/T
1486/1
22574 MNI
per sample
1485/1
Taxon
Atomaria sp.
1
9
13
N/A
1
in rotting hay and straw, as
well as in stable manure and
compost heaps, also in rotting
vegetation, beet heaps, fresh
grass cuttings, game food
debris, dry dung and wild
animal droppings, more rarely
in woodland litter, detritus,
grass tussocks and flood
debris, as well as at sap flows
on trees (Koch 1989b)
16
8
Ephistemus
globulus (Payk.)
mycetophagous, in mould
fungi; synanthropic, in barns,
etc, in houses on damp walls,
in granaries and warehouses,
in rotting provisions, corn etc.
In the wild, among mouldy
leaves, vegetation, near fungi
and tree fungi, Polyporus sp.
(Horion 1961)
1
1
14
Lathridius
minutus (grp.)
(L.)
Enicmus sp.
1
mycetophagous
Corticaria sp.
5
1
mycetophagous
90
Ecology
1469/T
22490
MNI
per
sample
1469/1
1486/T
1486/1
22574 MNI
per sample
1485/1
Taxon
Corticaria/
Corticarina sp.
2
mycetophagous
1
6
10
4
Aglenus
brunneus (Gyll.)
Lyctus linearis
(Goeze)
1
ancient broad-leaved woodland, also in timber yards and
in buildings, develops in dead
wood, especially of oak, beech
and ash, often in fresh oak
palings
(Hyman
1992);
especially in dry dead wood of
structural timbers (Robinson
1991)
1
1
4
Anobium
punctatum
(Deg.)
3
mouldering wood, in fungi
(Kenward 1975b);
barns,
stables, cellars and garden
centres, also field margins and
weedy places; under mouldy
straw, hay, chaff, straw
manure and mouldy chaff,
under dry mouldy manure in
chicken and dove cotes, in the
ground under rotting boards
in beet waste, single with
Talpa sp. and Microtus sp.
(Koch 1989a)
91
in furniture and tools; in
joinery timbers and flooring
and structural timbers of
buildings. Attacks willow,
alder and birch soon after the
timber is dry, but softwoods
require time to mature about
20 years and oak above 60
years before flight holes
appear (Buck 1958)
Ecology
1469/T
22490
MNI
per
sample
1469/1
1486/1
1486/T
22574 MNI
per sample
1485/1
Taxon
Ptilinus
pectinicornis
(L.)
2
in wood in debarked areas of
dry standing and fallen trunks
of hard deciduous wood,
especially in Fagus sylvatica,
but also Quercus, Carpinus,
Acer,
Ulmus,
Populus,
occasionally also in stumps
and thick branches; sometimes
in furniture (Koch 1989a)
Ptinus fur (L.)
1
2
2
common in mouldy straw and
hay in barns and heaps, in
cereal debris, in nests of
Passer domesticus, in carrion
and nesting materials in
pigeon lofts, sometimes in old
beehives, wasp nests and in
damp walls in lavatories, in
wild game food residues, in
twig heaps and in wood mould
of
hollow
trees.
Very
polyphagous (Koch 1989b).
Anthicus
formicarius
(Goeze)
1
in vegetable refuse, haystack
bottoms, compost heaps, etc.
(Buck 1954); syn. Omonadus
formicarius Goeze
92
Ecology
1469/T
22490
MNI
per
sample
1469/1
1486/1
1486/T
22574 MNI
per sample
1485/1
Taxon
Anthicus
floralis/
formicarius
(L.)/(Goeze)
1
A. floralis: in rotting and
mouldy hay and straw in ricks,
barns and stable manure
heaps; in rotting vegetation,
also fungi and compost; in
wood shavings; occasionally
on low plants as well as leaves
and twigs (Koch 1989a), syn.
Omonadus floralis L.; A.
formicarius: in vegetable
refuse, haystack bottoms,
compost heaps, etc. (Buck
1954)
1
decaying vegetation
1
non-obligate
synanthrope;
stored product pest
1
(s.l.)
1
Anthicus
sp.
in cellars, stalls, corn stores
and mills, more rarely in
deciduous woodland, parks
and gardens. Above all in
cereals and their products,
also in pigeon lofts and nests
of
Passer
domesticus,
occasion-ally
in
hollow
deciduous trees, mostly in
association with old bird
nests, under rotting bark, in
mouldy stumps (Koch 1989b)
Blaps sp.
scaber
in birds' nests, in hollow
trees, mostly owl and other
nests containing bones, and
in detritus of animal origin
(Jessop 1986)
1
1
Trox
(L.)
1
1
Tenebrio
obscurus (F.)
93
Ecology
1469/T
22490
MNI
per
sample
1469/1
1486/1
1486/T
22574
MNI
per sample
1485/1
Taxon
Aphodius
prodromus
(Brahm)
1
in all dung, especially horse
and human faeces; more
rarely in rotting vegetation,
also fungi, compost, stable
manure heaps, as well as in
grape debris and rotting fruit
(Koch 1989a)
2
Melolonthinae
indet.
N/A
1
2
Aphodius sp.
N/A
Phymatodes alni
(L.)
1
recently dead or decaying
Quercus spp. (Alexander
1994); also found in Alnus sp.
(Bullock 1993)
1
Chrysome-lidae
indet.
on Cruciferae, often on
disturbed or cultivated ground
and a pest of cultivated turnip,
Brassica sp. (Duff 1993)
1
1
Phyllotreta
nemorum (L.)
N/A
Phyllotreta sp.
1
found on vegetation in
disturbed and arable ground
Longitarsus sp.
2
N/A
1
Altica sp.
94
N/A
Ecology
1469/T
22490
MNI
per
sample
1469/1
1486/T
1486/1
22574
MNI
per sample
1485/1
Taxon
Scolytidae indet.
1
N/A
Curculionidae
indet.
1
N/A
Apion (Exapion)
difficile (Hbst.)
2
phytophagous associated with
Genista spp., possibly only on
dyer's greenweed, G. tinctoria
(Hyman 1992)
Apion (s.l.) sp.
1
associated with vegetation in
disturbed/arable land
Phyllobius sp.
1
N/A
Notaris
acridulus (L.)
1
larvae on roots of aquatic
grasses - Glyceria aquatica;
adults on Glyceria ssp. and
Polygonum
amphibium
(Hoffman
1958);
tall,
waterside vegetation (Duff
1993)
Hypera sp.
1
N/A
95
Ecology
1469/T
22490
MNI
per
sample
1469/1
1486/1
1486/T
22574
MNI
per sample
1485/1
Taxon
Micrelus ericae
(Gyll.)
1
on heaths, especially closed
Calluna heath, light pine
plantations, bogs, also dry
turf. Oligophagous on Calluna
vulgaris and Erica tetralix
(Koch 1992)
1
1
Ceuthorhynchus
contractus
(Marsham)
Ceuthorhynchus
(s.l.) sp.
polyphagous, especially on
Brassicaceae, but also on
Resedaceae
and
Papaveraceae,
in
winter
occasionally in grass tussocks,
hay, straw, leaves, twigs, moss
on trunks and flood debris.
Larvae in leaf mines (Koch
1992)
2
N/A
Ceuthorhynchinae indet.
1
N/A
Anoplura
Pediculus
humanus (L.)
1
parasitic; the human louse
Mallophaga
Damalinia ovis
(L.)
1
1
parasitic; the sheep louse
(Henriksen 1937)
96
Ecology
1469/T
22490
MNI
per
sample
1469/1
1486/T
1486/1
22574
MNI
per sample
1485/1
Taxon
Diptera
6
15
18
1
16
Diptera indet.
the sheep ked; lives in wool of
sheep
1
1
Melophagus
ovinus (L.)
Siphonaptera
Siphonaptera
indet.
1
Parasitic
1
irritans
the human flea; also found on
goats, pigs, badgers, and foxes
1000
Pulex
(L.)
anthophiles; the honey bee
20
mellifera
50
Apis
(L.)
50
Hymenoptera
15
Apoidea sp.
anthophiles
References for species’ habitats and diets cited in Buckland and Buckland 2006
97
Figure 4.8 % Number of species per ecological group
100%
Bees
90%
Ectoparasites
Structural Timbers
% Number of Species
80%
Synanthropes
Lathridiidae
70%
60%
Carcasses
Dung/ foul organic
50%
Disturbed ground/ arable
Marsh/ aquatic plants
40%
30%
Wood and trees
Meadowland
20%
Pasture/dung
Aquatic
10%
0%
Figure 4.9 % Minimum number of individuals reported for each species group
% MNI Coleoptera
100%
90%
Structural Timbers
80%
Synanthropes
70%
Lathridiidae
Carcasses
60%
Dung/ foul organic
50%
Disturbed ground/arable
Marsh/ aquatic plants
40%
Wood and trees
30%
20%
Meadowland
Pasture/dung
10%
Aquatic
0%
4.3.4 Palaeoenvironmental Reconstruction of Anglo-Scandinavian Coppergate
The aquatic and waterside environment
Coleoptera associated with aquatic (Group 1) and marshland (Group 5) habitats did not form a
high proportion of the assemblage. Although combined the number of species from the two groups
comprised 17 % of the total Coleoptera in the assemblage, the number of individuals recovered only
represented about 8 % of the total beetles. Several of these species could have lived around the river.
Trechus micros, in particular, is commonly recorded on the banks of running water (Lindroth 1974),
98
and Bembidion biguttatum has been found on the banks of rivers and ponds (Lindroth 1985).
Pterostichus melanarius and Bembidion gilvipes have been noted in flood debris and wash zones
(Koch 1989a). Moreover, a number of the Staphylinidae (though not restrictively so) occur in muddy
banks near water, e.g. Carpelimus bilineatus and Platystethus nitens. The majority of the waterside
and aquatic species most likely entered the archaeological context through association with the nearby
Figure 4.10 MNI by ecological group for 16-22 Coppergate entomofauna
1135
1150
1100
1050
1000
950
900
850
Aquatic
800
Pasture/ dung
Minimum Number of Individuals
750
700
Meadowland
Wood and trees
650
Marsh/ aquatic plants
600
Disturbed ground/ arable
Dung/ foul organic
550
500
Carcasses
Lathridiidae
450
Synanthropes
400
Structural Timbers
Ectoparasites
350
Bees
300
250
200
150
78
100
50
41 48
8
9 17 6 12 6
8
12
7
0
Species Ecological Group
rivers. While the fauna may have occupied similar micro-habitats to their ‘indigenous’ waterside
environment within Tenement C, i.e. puddles, mud, compost heaps, or damp decaying vegetation, the
relatively low number of individuals recovered for each species is not unreasonable given the site’s
99
proximity to the rivers. The beetles could have easily entered the structure through flight or, postmortem, blown in by the wind.
The woodland and scrub
The samples analysed from contexts 22574 and 22490 yielded a limited number of beetles
associated with wood and trees. 5.9 % of the coleopteran species and 2.4 % of the individuals from
the assemblage are wood dependent. These figures do not include the data for Anobium punctatum as
the species is almost entirely synanthropic. While Lyctus linearis is also strongly associated with
structural timbers, the beetle has been recorded in ancient woodland where it develops in dead wood
(Hyman 1992). The recovery of Phymatodes alni may indicate the presence of Quercus spp. (oak),
and the species typically inhabits wood with the bark still intact. The beetle Ptilinus pectinicornis is
especially associated with Fagus sylvatica (European beech); unlike Phymatodes alni, Ptilinus
pectinicornis prefers to infest debarked wood. Melanotus sp. is indicative of rotting wood and bark.
The wood entomofauna recorded from the samples is primarily suggestive of the presence of dead
wood rather than living trees and shrubs. Moreover, the samples did not yield any phytophagous
species, ground beetles, or staphylinids that are common in woodland communities. The absence of
non-wood associated woodland species implies that the ancient environment surrounding the 16-22
Coppergate site was not heavily forested. The wood-related species were probably transported to the
site with wood intended for use as timbers in structures, brushwood, and/or firewood.
Grassland, arable, and the open environments
Coleopteran species associated with open environments were fairly well represented in the
Coppergate assemblages, though not abundant. Meadowland fauna comprised 11.7 % of the species
and 6.9 % of the individuals recovered. The open environment-indicator species at the site were
primarily phytophagous or phytodetriticolous. Species that are considered to be specifically grass-root
feeders, i.e. assigned to Species Group 11, were not present. The samples yielded coleopterous
species, such as Tachinus rufipes, which have been recorded at grass-roots (Duff 1993) but are more
common in decaying grasses.
Some of the phytophagous Coleoptera recovered from the Coppergate samples are fairly hostspecific, including:
Apion (Exapion) difficile
Genista tinctoria
Micrelus ericae
Calluna vulgaris and Erica tetralix
Brachypterus sp.
Urtica sp.
The weevil Apion difficile is highly indicative of the presence of the low-lying sub-shrub
Genista tinctoria, dyer’s greenweed. G. tinctoria performs poorly in wet soils and is believed to
favour dry meadowland and heathland environments (Bown 1995). The plant is not likely to have
100
been indigenous to the riverside environment near the Coppergate. The two individuals of A. difficile
present in the samples were likely transported to the site along with the greenweed, which may have
served as a dye source.
The weevil Micrelus ericae is supportive of a heathland connection. M. ericae feeds on
Calluna vulgaris (common heather or ling heather) and Erica tetralix (the cross-leaved heath or bell
heather). Both plant species inhabit wet heathland and bogs. Today, ling heather and to lesser extent
bell heather are ‘coveted’ sources of honey in Britain, and beekeepers will establish apiaries near
heather stances (Weightman n.d.). Moreover, Beekman and Ratnieks (2000) found that honeybees
would forage Calluna vulgaris stances located over 9.5 km from their hive. While the AngloScandinavians may have also been exploiting heather for honey, especially given the large number of
honeybees recovered from the site, the presence of M. ericae suggests the physical transportation of
the plant to the site.
The British Brachypterus species are typically monophagous on nettles, e.g. B. urticae F. and
B. glaber Steph. The nettles may have grown along the river bank. However, the disturbed ground of
a grassland environment or woodland margin would also support the plants. The beetle was present
but not abundant in the assemblage.
7 % of the Coleoptera species and 2.5 % of the minimum number of individuals were
associated with arable or bare ground. Two individuals of Phyllotreta nemorum were recovered,
which as mentioned above, is associated with Cruciferae. Many of the Ceuthorhynchinae species feed
on weeds of the family Cruciferae. While common in flood plains and wash zones, the carabid
Pterostichus melanarius also occurs in high numbers on the bare ground between ground level plant
stems as well as in low numbers in grasslands (Robinson 1979).
There was little evidence of pastureland species recovered from the site. A large number of
Aphodius species are associated with dung in the field. A. prodromus is most commonly recorded in
horse dung but has been noted in decaying vegetation (Koch 1989a). Cercyon atricapillus is also an
indicator of horse dung usually in pastures. Unfortunately the majority of the dung-related beetles,
which were recovered from the site, are difficult to interpret because they are also known to inhabit
decaying vegetation. Although the samples contained no purely dung related fauna, which makes the
presence of a pastureland-herbivore connection difficult to discern, the Coppergate contexts produced
a number of insect species associated with decaying vegetation. Furthermore, the samples contained
decomposer species like Xantholinus linearis, which appear to avoid faeces altogether (Buck 1955;
Robinson 1979). If dung was present at the site, it was not abundant and may have been kept in low
quantities to fuel fires.
Beetles associated with grassland, arable land, and pastureland environments suggest an
imported component rather than local signal to the site. While Brachypterus sp. may be connected to
the local riverside environment and Phyllotreta nemorum may have arguably inhabited the vegetation
101
in the local gardens, Apion (Exapion) difficile and Micrelus ericae are representative of plants that
may have been transported from the hinterland regions such as the heathland on Vale of York.
Other habitats
Several of the insect species recovered from the Coppergate samples are synanthropic. Beetles
especially associated with structural timbers comprised 4.8 % of the coleopteran individuals. Anobium
punctatum was present in four of the five samples. The species is strongly associated with worked
wood and may infest furniture, structural timbers, and flooring as well as the wood in tools (Buck
1958). Although Lyctus linearis can occupy a similar habitat as A. punctatum, it was only present in
one of the samples. The beetle Ptilinus pectinicornis was also present and has been known to infest
furniture wood (Koch 1989a).
Many of the decomposer beetles are associated with decaying hay or straw and can be
categorised as what Carrott and Kenward (2001; Kenward and Carrott 2006) called house fauna.
Decomposers considered house fauna are typically associated with dry, possibly mouldy habitats.
Ephistemus globulus, Xylodromus concinnus, and Acritus nigricornis were ranked present to abundant
in the samples and are common in decaying hay and straw. In sample 1487, Hall and Kenward
(1999b) calculated the MNI of the dry decomposers as 35 %. Crataraea suturalis is also associated
with mouldy straw and is strongly synanthropic (Harde 1984).
Moreover, the mycetophagous
Lathridiidae fauna from Species Group 8 and Cryptophagus scutellatus are frequently recorded in
mouldy hay in barns. The hay indicator species suggest the presence of hay and straw at the site,
which may have served as floor litter or thatch.
Ptinus fur is also representative of Carrott and Kenward’s (2001) house fauna group. Although
the beetle has been found living in mouldy hay and straw, it is rather polyphagous and has been noted
in birds’ nests and old beehives (Koch 1989b).
While potentially associated with the material
comprising the litter and/or thatch, the P. fur individuals may have also inhabited the hives of
associated with the abundant remains of honeybees, MNI 1135, which constituted 74.5 % of the
assemblage. While the bees may have been transported with the hives to the site and then killed off
during the honey processing, the number of individuals suggests that the hives were most likely being
kept on the site. This is further supported by the recovery of large numbers of bees from for of the
five samples, which implies that Apis mellifera were fairly prevalent around the site as the mass death
assemblage was not isolated to a single location.
The beetle Ptinus fur has also been considered a pest of stored cereal products (Mound 1989).
Tenebrio obscurus and Blaps sp. are present in low numbers and support the presence of stored
products, such as flour or cereals. However, T. obscurus has also been noted in rotting bark and bird’s
nests (Koch 1989a), and Blaps sp. feed on a range of vegetable matter including straw waste (Horion
1956). While grains and cereal products may have been kept at the site, the recorded insect fauna is
102
not conclusive as it neither proves nor disproves the presence of cereals in the contexts. The more
indicative grain species, i.e. Sitophilus granarius, Cryptolestes ferrugineus, and Palorus ratzeburgi,
were not present in the assemblage but also lack any convincing archaeoentomological record to
support their presence in Britain between the end of the Roman occupation and the Norman Conquest.
Beetle species that are typically representative of animal carcasses and carrion comprised 4.7 %
of the Coleoptera. Omosita species tend to infest carrion in the later stages of decay (Robinson 1991).
Beetle species of the genus Dermestes have been known to feed on dead insects, animal carcasses,
stored meats, skins, hides, furs and bones (cf. Hinton 1945). Two individuals of the beetle Trox
scaber were also present in the contexts. The beetle has been noted in bird’s nests, especially those
containing bones and animal remains (Jessop 1986). However, Koch (1989a) noted T. scaber on horn
cores, fleeces, animal skins, bird carcasses, wasp nests, and mouldy straw.
The identified
carrion/carcasses associated fauna recovered from the site tend to be more generalist feeders rather
than being strongly indicative of a specific resource or material.
The remains of ectoparasites were found in the Coppergate samples.
The sheep ked
Melophagus ovinus and the louse Damalinia ovis are associated with sheep. Given the paucity of
dung beetles in the contexts, the site was probably not used to keep sheep. However, both M. ovinus
and D. ovis have been used by researchers as evidence of wool processing (Buckland and Perry 1989;
Jaques et al. 2001; King 2006). The human flea Pulex irritans and the human louse Pediculus
humanus were also present at the site.
4.3.5 Palaeoclimatic Reconstruction
TMinHi
26
18
18
-9
-8
-7
11
15
28
18
-23
-7
13
7
7
0
10
7
9
11
11
27
24
22
8
11
39
22
NSPECIES
TMinLo
15
15
15
TRange
Lo
TRange
Hi
TMaxHi
Sample
22574.1485/1
22574.1486/1
22574.1486/T
22490.1469/1
22490.1469/T
Site
TMaxLo
Table 4.4 MCR estimates for fill contexts from Period 4b Tenement C
7
13
6
0
1
20
Overlap
85.71429
92.30769
83.33334
0
100
95
(calculated using Buckland and Buckland 2006)
The palaeoclimate predictions calculated using BugsMCR program (Buckland and Buckland
2006) are shown in Table 4.4 and Figure 4.11. As with the 7-15 Spurriergate samples, only the
carnivorous and scavenging beetle species were assessed. Twenty species were analysed [Appendix
1C] and tabulated the temperature of the warmest month as ranging between 15 ˚C and 18 ˚C and the
103
temperature for the coldest month between -7 ˚C and 7 ˚C. The MCR prediction for the Period 4b,
16-22 Coppergate Tenement C samples indicates slightly warmer temperatures during the coldest
month relative to the 1st century AD York, but the insect remains from both periods suggest a similar
range of temperatures during the warmest month. The range of temperatures estimated for Period 4b
16-22 Coppergate Tenement C samples approximates the range evidenced by beetles from other
Anglo-Scandinavian sites in York [Table 4.5].
4.3.6 Discussion: The Environment and Climate of Period 4b 16-22 Coppergate and Implications
for Culture Contact
Through the archaeological excavations conducted at 16-22 Coppergate, the insect remains
recovered from contexts 22574 and 22490 were believed to be associated with the inside of a
Figure 21 16-22 Coppergate, York MCR estimates by sample
30
25
TMax
20
15
10
5
22490_1469_/T
22490_1469_/1
22574_1486_/T
22574_1486_/1
15
10
5
0
-5
-10
-15
-20
-25
22574_1485_/1
TMin
0
(calculated using Buckland and Buckland 2006)
104
TMaxHi
TMinLo
TMinHi
TRangeLo
TRangeHi
N-SPECIES
Overlap
1-9 Micklegate
15
18
-7
2
16
22
26
100
6-8 Pavement
15
18
-7
6
12
22
48
100
5-7 Coppergate
15
18
-7
6
11
22
35
100
Environmental
Report
TMaxLo
Table 4.5 MCR estimates for Anglo-Scandinavian sites in York, UK
Sample
Period 3, 16-22
Coppergate
Period 4a, 1622 Coppergate
Period 4b, 1622 Coppergate;
entire site
16
15
18
18
-6
-7
6
6
11
11
22
22
48
23
100
100
Kenward and
Hall 2000
Hall et al. 1983
Hall et al. 1983
Hall and
Kenward 1999a
Hall and
Kenward 1999b
Hall and
Kenward 1999b
16
18
-6
6
11
22
40
100
(calculated using Buckland and Buckland 2006)
structure, i.e. Tenement C. However, even without the aid of the archaeological interpretations, the
ecological assessment of the insect remains suggests the presence of a structure [Figure 4.13].
Anobium punctatum and other Group 10 species implying the presence of worked wood. The samples
yielded a number of dry decomposers and hay-associates which are typical of thatch and floor litter.
Moreover, the contexts did not contain an abundance of outdoor fauna. While a few riverside and
aquatic species were present in the samples, their presence was minimal and in the expected range
given the proximity of the River Foss and River Ouse. Kenward and Allison (1994b) include aquatics
and waterside species as well as decomposers and wood associates [Figure 4.14] in their list the
original habitats of urban sites. The majority of the other outdoor species, such as those associated
with meadowland, pastureland and heathland, are indicative of animal and plant materials, which are
reflective of two potential industries: 1. honey and wax and 2. wool and dye.
The recovery of large quantities of honeybee remains suggests that the Anglo-Scandinavians
were exploiting Apis mellifera. The physical presence of honeybee remains indicates that the AngloScandinavians would have had access to ‘local’ honey and wax resources rather than needing to
import the commodities.
The number of individuals in the death assemblage suggests that the
honeybees would have kept a hive near, if not within, the structure. Furthermore, although largely
105
Figure 4.13 Likely sources of insect remains in building deposits (Kenward 1985, 105)
Key:
Broad arrowsautochthones
Narrow arrowsallochthones
Figure 4.14 Probable starting habitats for urban insects
Corpses
Nests and Tree
Hollows
Dung
Urban
Site
Dead Wood
Plant Litter
Waterside
Subterranean
(King 2006, 79; Reconstructed from Kenward and Allison 1994b, 60-61)
hypothetical and at risk of stretching the evidence, the presence of heather at the site, evidenced by
Micrelus ericae, may have been gathered from the heathland, and potentially the turnip Brassica sp.,
suggested by Phyllotreta nemorum, from the meadowlands (see Columella’s RR IX 1941) to
encourage the pollination of sweet flowers and plants. The alternative is that the hive was located
elsewhere and the bees and the hive were transported to the site for processing, but individuals would
have likely been lost during transport resulting in a smaller death assemblage. However, if numerous
hives were being transported and processed, a large death assemblage would also be expected under
good preservational conditions. While the first possibility seems most likely, the interpretation is
purely speculative and the possibility of hive transportation should not be dismissed. Given the
documentary evidence describing the handling and relocation of beehives from the Roman era (cf.
Columella’s RR IX 1941), it is likely that Anglo-Scandinavians would have been familiar with
methods of transporting active hives.
Did the Anglo-Scandinavians import the honeybee to Britain? There are several races of
honeybees, and Kenward (pers. comm) has identified the Coppergate remains as A. mellifera mellifera
L., the European Dark Bee, which is a cold-tolerant species. Today, natural distributions of the
European Dark Bee are found as far north as 60˚ N latitude (Ruttner et al. 1990). Although the
honeybee is capable of generating and regulating heat within clusters (Ruttner et al. 1990), Villa and
Rinderer (1993) reported the death of whole colonies when temperatures were held at 0˚ C for 10 days.
Considering the estimated coldest month temperature range of -7 ˚C and 7 ˚C in Anglo-Scandinavian
York, colonies of Apis mellifera mellifera would have been able to survive in the wild. As such, the
palaeoecological assessment cannot provide insight into whether the Anglo-Scandinavians imported
hives of honeybees from afar.
The insect remains recovered from the Tenement C contexts are also indicative of the
processing of wool and dyeing. As mentioned above, there was little conclusive evidence to support
the presence of dung and very little pastureland fauna at the site, which implies that the sheepassociated ectoparasites, i.e. Melophagus ovinus and Damalinia ovis, do not indicate the stabling of
sheep but rather the processing of wool.
processing at the site.
Several other faunas support occurrence of the wool
The carrion and carcass-associated species, especially Trox scaber and
Dermestes sp., would have been attracted to the stored wool and skins. The foul organic species from
Species Group 7 may have been attracted by storage and use of urine to bleach and delouse the wool
(Stead 1982; Buckland and Perry 1989; King 2006). The recovery of Apion (Exapion) difficile
evidences the presence of the plant dyer’s greenweed, Genista tinctoria. G. tinctoria and quite
possibly the heather species, Calluna vulgaris and Erica tetralix, could have been employed as dye
plants (Kenward and Hall 1995; Hall and Kenward 1999b). The dye plants may have been brought to
the site to aid in the dyeing of the wool or other fabrics.
In an effort to determine whether the wool was acquired from local or imported sheep, the
thermal requirements of the ectoparasites were considered. Graham and Taylor (1941) reported no
emergence of Melophagus ovinus from puparia at temperatures below 18 ˚C and the death of newly
107
emerged keds within 24 days at 4 ˚C when the species was separated from its host. Damalinia ovis
appears to be less cold tolerant. Murray (1957ab) demonstrated that temperatures between 32 and 40
˚C were required for the sheep louse to lay eggs, and Murray (1960b) showed that morphogenesis of
the eggs was only completed within the temperature range 30-39 ˚C. However, when development
was completed, the eggs hatched between 22 and 42.5 ˚C. It is important to note that temperatures
near the skin of the sheep are warmer than the ambient temperatures. Murray (1960a) reported that
temperatures near the skin, which was covered by five centimeters of wool, reached 39 ˚C when the
atmospheric temperature was 28 ˚C.
The temperature requirements of the ectoparasites imply that neither species was likely to have
evolved in Britain. However, M. ovinus would have been able to survive and reproduce in the wool
microclimates once introduced to the region. As a wingless fly, the species is believed to spread only
through direct contact (Small 2005). While the eggs of Damalinia ovis would have been able to hatch
when kept in the wool, the thermal requirements necessary for ovipositioning to occur and the
morphogenesis of the eggs would have barely been attainable during the warmest months in AngloScandinavian York. Considering an 11˚ C temperature increase from the skin level microclimate to
atmospheric temperature, a TMAX of 18˚ C would suggest a temperature under the wool
approximating 29˚ C. If the sheep were kept in sheltered environments such as stables, the skin-level
temperature would have been even higher. Moreover, if the Anglo-Scandinavians were keeping sheep
species with longer hairs or thicker wools, the skin-level temperatures would have been warmer,
which would have theoretically enabled the sheep louse to reproduce and survive in colder
environments. Regardless, the presence of D. ovis implies a connection to warmer climates, which
would have allowed the louse to enter Britain.
Unfortunately, temperature requirements and tolerances of the probable dye plant associates,
Apion (Exapion) difficile or Micrelus ericae, are not known.
According to modern distribution
regards, both weevils appear to be fairly cold tolerant. A. difficile has been reported as far north as
54.1˚ N in the United Kingdom (GBIF 2009a) and has been noted in Dorset, Sussex, Kent and
Gloucester in England (Morris 1990). Morris (1990) also reports the distribution of the beetle
extending north to Denmark. M. ericae is a subarctic to temperate species (Böcher 1995) and has been
reported as far north as 64.7˚ N (GBIF 2009b) and is distributed throughout central Europe (Koch
1992) and the United Kingdom (Joy 1932; Duff 1993). The modern distribution of both species
suggests that they may have been indigenous to Britain. However, neither species is likely to have
been autochthonous to the Coppergate site given their habitat preferences (see above), and both were
probably transported to the site from York’s hinterlands.
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4.3.7 Summary
The samples examined from Period 4b 16-22 Coppergate, York contained several indicator
species associated with human commodities and industries. Anobium punctatum and Lyctus linearis
indicated the presence of structural timbers, which were most likely associated with the post-andwattle building. The recovery of Apion (Exapion) difficile suggested that dyer’s greenweed was stored
and/or used at the site. A similar interpretation was offered for the heathland species Micrelus ericae
and its association with heather. The Tenebrio obscurus and Blaps sp. specimens may indicate the
presence of stored cereals; however, as the species have also been associated with other habitats
including nests, straw waste, and bark, which may have been present at the site, the findings were
inconclusive. Wool processing was occurring at or near the site as evidenced by the sheep ked
Melophagus ovinus and the sheep louse Damalinia ovis. Trox scaber may also support the processing
and/or storing of animal materials such as wools, hides, and furs. The large number of honeybees
reported from the contexts evidences the availability of honey and wax to Anglo-Scandinavians.
The comparison of the species’ thermal requirements and the site’s MCR revealed that most of
the indicator species would have been able to survive away from human habitation in AngloScandinavian Britain. D. ovis was an exception as the species’ preferred temperature range was
indicative of warmer climates. However, the sheep louse may have been able to complete its life cycle
once entering Britain if it infested sheep with thick or longhaired wool and/or sheep which were kept,
at least part of the time, in indoor environments, e.g. stables.
Several of the indicator species are strongly associated with environments, e.g.
heathland/moorland, which would not have likely been available in urban York. The species are more
indicative of environments that, at least today, may be found in the York’s hinterland regions, such as
the Vale of York or Askham Bog. The presence of Micrelus ericae and Apion (Exapion) difficile
suggest that the Anglo-Scandinavians were exploiting the hinterland environments and having Genista
tinctoria, Calluna vulgaris, and/or Erica tetralix transported to the site. However, it is possible that
both the plants and their associated beetles may have been imported from a greater distance, but such
resolution is beyond the scope of a palaeoecological assessment.
4.4 Conclusion
The palaeoecological approach provides a very versatile tool for the assessment of
archaeologically recovered insect remains.
Insect subfossils have been effectively employed as
bioindicators of palaeoecosystems and archaeological reconstructions as a result of their ecological
diversity, their tendency to be often ignored or perceived as unimportant to humans, and their
sensitivity and rapid reaction to environmental change (see Coope 1977b; Elias 1994; Bain 1997;
1998; Kenward 1999; Whitehouse 2006). Insects, therefore, have huge potential to stand as evidence
109
of past human activity, living conditions, diet, climate and ecology. While archaeological insects pose
questions about the origin and development of regional faunas, palaeoentomological studies have
revolutionized perceptions about rates of evolution and the morphological and ecological constancy of
species, as well as climatic change. The fossil remains of insects buried deep in the earth are a crucial
but often neglected part of investigations of the human, and wider ecological and climatic, past. In
regards to the present evaluation, the analysis of archaeoentomological subfossils enables researchers
to glean a better understanding of palaeoeconomic activities.
By superimposing the habitats and ecological requirements of modern insect species over the
fossil record [Chapter 2], certain insect remains retrieved from the archaeological sites are invaluable
as indicators of the presence of specific plant and animal materials, which may have been exploited by
humans in the past. The insects may stand as primary, e.g. Apis mellifera indicating the availability of
honey and beeswax, or secondary evidence, e.g. Sitophilus granarius suggesting the presence of stored
grains, of exploitable commodities.
Although insect remains may be ‘reliably’ used as indicators of materials, they have limited
effectiveness as a tool for discerning culture contact, human movement, and palaeoeconomics. By
assigning species to broad ecological groups, the presence of certain environments is discernable, e.g.
grassland, pastureland, heathland, woodland, etc. Moreover, quantitative assessment of the material
may be employed to determine the autochthonous and allochthonous components (see Kenward 1978;
Perry et al. 1985). Unfortunately, palaeoenvironmental tools do not provide a means of assessing
distance and researchers are forced to speculate as to the most likely source of the allochthonous
species. For example, Apion (Exapion) difficile indicates a heathland or meadowland connection and
the presence of Genista tinctoria in Anglo-Scandinavian York, but where did the dyer’s greenweed
originate? Today, G. tinctoria may be found in Vale of York (A. Hall pers. com); however, it is also
has a continental European distribution ranging from Spain in the south to Norway in the north (see
GRIN 2009). Were the Anglo-Scandinavians electing to import the dyer’s greenweed, or were they
harvesting it from the local hinterlands?
Another major limitation of single site palaeoecological assessments is that it does not provide
a timeframe for the initial introduction of potential exploitable materials. For example, was G.
tinctoria available in Northern England prior to the arrival of the Anglo-Scandinavians, or do the
modern populations stem from individuals purposely introduced in the past?
This is especially
apparent in the palaeoclimatic aspect of the present analyses. While a comparison of the MCR data
and species’ temperature requirements may help identify probable ecological outlier taxa, it does not
account for the possibility that the species may have been introduced during earlier periods.
A timeframe for the introduction of various insect species may be established through
application of methods which consider the presence of individual species at multiple sites and assess
changes in spatial and temporal changes in species’ distribution, e.g. the biogeographical approach
110
[see Chapter 5]. Furthermore, isotopic and genetic analyses may be of assistance in helping to
overcome the problem of multi-region resource availability by identifying geographic and
phylogenetic similarities and differences within and between assemblages [see Chapters 6, 7, and 8].
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Chapter 5
Grain Pests: An Archaeobiogeographical Account of the History of their Dispersal
112
5.1 Introduction
Grain and other storage pests cause significant depletion of human food resources at the
present day (e.g. Tyler and Boxall 1984; McFarlane 1989; Payne 2002), and the beetles are among the
most economically important. Documentary records show that in the 19th and early 20th century they
represented a very serious cause of food loss (cf. Munro 1966), and remains recovered from
archaeological deposits indicate that grain beetles were often abundant at earlier dates as well. These
beetles, which are strongly synanthropic in most of their range, are of substantial interest from the
ecological point of view, as aliens which have invaded artificial habitats, often alongside native
species. From a biogeographical angle they are significant as species spread by human activity, often
well beyond their naturally viable distributions. Where did they originate, and when did they spread?
They are also of considerable significance in studies of early agriculture: when did the first farmers
encounter these pests, and how significant were they among the tribulations Neolithic people endured?
Biogeographers typically recognise three distinct types of dispersal pathway by which
organisms may spread between areas through natural means: the corridor, the filter, and the
sweepstakes route. In the corridor route, the pathway may include a wide variety of habitats with the
areas at the two ends possessing an almost identical biota, e.g. pre-Ice Age Eurasia. The corridor
pathway would enable the majority of organisms to transverse between the two end areas with little
difficulty. The filter pathway consists of a more limited variety of habitats so that only organisms that
can exist in those habitats can disperse between the interconnecting regions, e.g. the tropical lowlands
of Central America. In the third type of dispersal pathway, the end regions are isolated as the result of
the interconnecting regions consisting of completely different environments, e.g. islands surrounded
by sea (Cox and Moore 2000).
In The Ecology of Invasions by Animals and Plants, Elton (1958) introduced the concept of
man as an impetus for the passive distribution of animals and plants beyond the prescribed boundaries
of their ‘original’ geographic range.
Building on that premise, Buckland (1981) reviewed the
archaeoentomological records available at the time for stored product pests and attempted to trace their
dispersal. While Buckland successfully demonstrated that pests were capable of being transported by
man in the past, he was unable to conclusively deduce patterns of movement or origins for the
evaluated species due to a paucity of fossil, particularly non-British, data.
Buckland offered
speculations and pleaded for the undertaking of more archaeoentomological evaluations in Eurasia. In
this chapter, the currently available fossil and literary evidence is employed to attempt to take the story
forwards and to ascertain the geographic origins of the stored cereal insect pests, their mode and route
of dispersal in the past, and their likely impact on early societies from the Neolithic Period through the
Roman Era.
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5.2 The Grain Fauna
The beetle fauna associated with stored cereals consists of a group of species which have been
ecologically classified in the northwest European archaeological context by Kenward (1978; 1997) as
‘strong synanthropes’, mostly thermophilic and wholly dependent on artificial habitats for survival in
the region.
This does not imply that the individual species are unable to survive beyond the
boundaries of the human-created artificial environments in other regions, as the classification is
climate-dependent, although in fact some of the species are virtually unknown in natural habitats. A
brief review of the biology of the typical grain pests is needed in order to set the archaeological
records in an ecological context.
Stored grain insects are often grouped based on their capacity or ability to infest undamaged
kernels as primary or secondary pests. Primary pests are
1.) “capable of successfully attacking, feeding and multiplying on previously undamaged grains;
2.) are adapted to feed on a narrow range of commodities;
3.) usually cause very distinctive damage;
4.) usually develop within the grains, and often complete their entire development within a single
grain;
5.) are selective in their egg-laying behaviour;
6.) often infest the ripening crop before harvest; and
7.) usually cannot develop on the same food if the grains are ground (milled),
[and] secondary pests are
1.) not capable of attacking previously undamaged grains, but can only attack and breed in grains
that have been damaged by primary pests, physical damage by bad handling, threshing, drying
or intentional processing that removes or damages the seed coat;
2.) usually attack a very wide range of commodities;
3.) usually cause non-distinctive damage;
4.) sometimes develop within grains, but never complete their development within a single grain;
5.) do not usually have selective egg-laying behaviour;
6.) are very rarely found on the crop at harvest; and
7.) are usually capable of developing on the same food after it is ground” (Semple et al. 1992).
While a number of cereal pests have been recovered from non-synanthropic situations, mostly
in tropical and subtropical areas, the granary weevil, Sitophilus granarius (L.), has yet to be found in a
natural habitat and its origin is particularly uncertain. Hoffman (1954) has recorded the weevil from a
wide range of stored products including wheat, rye, barley, maize, oats, buckwheat, millet, chickpeas,
and even chestnuts, acorns and cornmeal; however, despite its catholic tastes, it is most common today
as a primary pest in stored cereal crops. Buckland (1990) has observed that the caryopses of wild
114
cereals may be too small for successful breeding as the larvae develop entirely inside the grain or seed,
and on-going experiments at the laboratories at the University of York appear to confirm that S.
granarius is unable to develop or unwilling to lay in grains smaller than millet. While both Zacher
(1938) and Howe (1965) have proposed acorns as the original primary host of the weevil, the species
seems unable to breech undamaged shells, and the micro-habitat similarities between rodent stores and
the grain stores of man point to rodent stores as a more likely origin.
The adult weevils are
characterised by a forward, snout-like extension of the head, the rostrum, which bears the mouth parts.
S. granarius has elongated oval punctures on the dorsal surface of the prothorax and has elytra that are
longitudinally grooved with fine punctures. In contrast with its congeners S. oryzae and S. zeamais,
which in the tropics may infest crops in the field (Krantz et al. 1978), the granary weevil is flightless,
and it is capable of adjusting to unheated indoor habitats in the cooler climates of Northern Europe.
As a poor disperser (cf. Mlambo 1980), its present cosmopolitan distribution can be attributed to
accidental transportation by man.
The rice weevil, Sitophilus oryzae (L.), differs morphologically from the granary weevil in the
deep round punctures on its prothorax and is almost identical to its sibling species the maize weevil,
Sitophilus zeamais (Mots.). While mating incompatibility confirmed separate species (Floyd and
Newsom 1959), only the aedeagal and eighth sternite of female characters appear consistent, as
determined by DNA analysis (Hidayat et al. 1996), for morphological distinction between S. oryzae
and S. zeamais. Both species are flighted and are believed to be of tropic origin. Despite their names,
they attack a wide range of stored cereals: rice, rye, maize, wheat, barley, millet, etc (Harde 1984), as
primary pests. The rice weevil has been known to infest crops in the field (Kranz et al. 1978; although
this has been contested by some researchers, e.g. Mlambo 1980) and has been recorded under bark and
on leaves in North America (Dillon and Dillon 1972). However, in Central Europe the species are not
found in the open but only in artificial environments (Harde 1984). Because of the two species’ tropic
preferences, their presence in temperate regions can be viewed as invasive rather than native.
The bostrychid Rhyzopertha dominica (F.), the lesser grain borer, is a primary pest of stored
grain as an adult and larvae. However, it has also been recorded on flour, dried tubers (Hill 1994a),
and seeds. While the lesser grain borer was originally identified and described from South America, it
is now completely pantropical (Munroe 1966) and appears to have had an Old World presence and
origin (Kislev 1991). The pale to dark reddish-brown species is recognisable by the transverse row of
teeth at the front of the pronotum (Kislev 1991). Like the Sitophilus weevils, R. dominica is capable
of causing great damage to food supplies through extensive feeding.
The saw-toothed grain beetle, Oryzaephilus surinamensis (L.), is considered a secondary pest,
typically feeding on grain previously attacked by primary pests such as Sitophilus granarius. The
adults have been noted to be carnivorous, feeding on larvae, but they also feed on meal and other
ground starchy foods as well as previously damaged or spoiled grains. Although the saw-toothed
115
grain beetle has been recorded in warehouses, mills, granaries, and brewery silos (Zacher 1927),
Hunter and others (1973) speculate that its primary habitat is under loose bark, and Horion (1960)
additionally suggest fungoid timber. Buckland (1990) puts forth that the fungus may be a necessary
intermediary between the natural and artificial environments however once established that the damp
grain could mimic the micro-habitat conditions of the under bark pabulum resulting in a suitable
environment for the species. O. surinamensis is recognisable by the six tooth-like projections along
each side of the prothorax. While capable of flight, the worldwide distribution of Oryzaephilus
surinamensis is most likely due to the transportation by man.
The flat grain beetle or rusty grain beetle, Cryptolestes ferrugineus (Steph.), is another
secondary pest of stored product cereals and is largely associated with processed foods. It is found in
wheat, ripe, maize, meal, and flour as well as dried fruits (Horion 1960).
Like Oryzaephilus
surinamensis, the original habitat of C. ferrugineus may have been under loose bark (Hunter et al.
1973) where it probably fed on fungi (Halsted 1993) and may have found a similar micro-habitat
amongst the stored grains. Cryptolestes ferrugineus is a very small (1.5 mm) reddish-brown flattened
beetle with long filiform antennae (Mound 1989). Although the flat grain beetle is cosmopolitan,
Halsted (1993) suggests that it is being replaced by its congener the Mediterranean flat beetle, C.
turcicus (Grouv.), in temperate regions.
As the two species have similar tastes, interspecific
competition may have impacted survival and distribution in the past.
Palorus ratzeburgi (Wiss.), the small-eyed flour beetle, is found in stored cereal products,
particularly mouldy grain residues previously attacked by grain weevils (Brendell 1975). The smalleyed flour beetle is known to be predacious on other pests as well as feed on the faeces of Sitophilus
granarius (Pals and Hakbijl 1992). Having been recorded living under bark in Germany (Reitter 1911
cited in Solomon and Adamson 1955), P. ratzeburgi possesses a similar ecology to C. ferrugineus and
O. surinamensis and mostly likely adapted to stored product environment in a similar fashion.
Palorus ratzeburgi is noted as a small (2-3 mm), oblong, flattened reddish-brown beetle. The sides of
its head are not strongly flexed upwards and the eyes are small and round (Brendell 1975). P.
ratzeburgi occupies a similar habitat to its congener P. subdepressus (Woll.), but while both are found
in the Mediterranean region, P. subdepressus is not established in Britain (Brendell 1975) and is
particularly widespread in the tropics (Rees 2007).
Tribolium castaneum (Hbst.), the rust-red flour beetle, is a secondary pest of stored grain and
an important pest of cereal products, flour and bran (Brendell 1975). It is characterised by its parallel
sides and, in adults, the eye partly divided by a side margin of the head (Kislev 1991). Although it is
now cosmopolitan, Hinton (1948) believes it to have originated in India where it has been recorded in
the wild. Outside of the tropics, it is largely restricted to heated buildings (Brendell 1975) and
occasionally under bark (Whitehead 1999). It appears unable to survive cold temperatures (Solomon
and Adamson 1955).
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The tenebrionid Tribolium confusum (Duval), the confused flour beetle, is more cold tolerant
than its congener T. castaneum and has been regarded as a temperate species (Solomon and Adamson
1955). Kislev (1991) states that it differs morphologically from the rust-red flour beetle by possessing
a wider separation between the eyes underneath the head. The confused flour beetle has been recorded
infesting stored grains and cereal derivations where it is believed to be a secondary pest (Brendell
1975). Although Tribolium confusum may be regarded as a temperate species, Panagiotakopulu
(2001) notes that, like T. madens (Charp.), it is thought to be of African origin.
While the species described above have an established, if not extensive, fossil record, the
archaeological presence for some of the other beetle grain pest fauna is more sporadic, but noteworthy.
The thermophilic khapra beetle, Trogoderma granarium (Everts), is a small (1.5-3.0 mm), oblong oval
insects with unicolorous elytra and evenly rounded eyes (Mound 1989). While the adults do not
normally feed, the khapra beetle larvae feed almost exclusively on grain and cereal products (Munro
1966). Tenebroides mauritanicus (L.), the cadelle, is a large (5-11 mm) shining-black oblong beetle
with a prothoraxic base distinctly separated from the base of the elytra (Mound 1989). As both adults
and larvae, the cadelle attack cereals, cereal products, nuts, and dried fruit and have been known to be
partly predacious (Dillon and Dillon 1972). The cadelle is larger than most stored product pests other
than the Blaps and Tenebrio species, which have a preference for grain but are largely omnivorous
scavengers. Similarly, the lesser mealworm beetle, Alphitobius diaperinus (Panz.), attacks a wide
range of stored products including grain but is primarily an omnivorous feeder (Brendell 1975).
Alphitophagus bifasciatus (Say), the two-banded fungus beetle, is commonly recovered from mouldy
and decaying grain and vegetable products in stables, granaries, and warehouses where both the adults
and larvae feed on mould (Brendell 1975). Some of the species usually listed among ‘pests’ in the
stored products literature are in fact quite eurytopic, able to exploit material such as birds’ nests and
debris under bark in nature, and thatch and litter in settlements. One such beetle is the biscuit beetle
(or drugstore beetle), Stegobium paniceum (L.), a pest of a range of farinaceous foods (Buck 1958). It
tolerates quite low temperatures.
5.3 Records of the Fauna
5.3.1 Pre-Roman Middle East
At the present, the Har-ra==Hubullu Tablets XI-XV contain the earliest written zoological
record supporting the human acknowledgement of the presence of stored product pests, listing 33
names of crop and stored product pests (Landsberger 1934). During the nineteenth century, these
cuneiform texts were recovered from the royal library of the Assyrian king Ashurbanipal (2618-2577
BP) at the ruins of Nineveh in Mesopotamia. While it was compiled during the ninth century B.C. in
117
bilingual Sumero-Akkadian script, the tablets are believed to be based upon Hammurabian period
(3678-3636 BP) lists which in turn developed from even older ones (Harpaz 1973).
On Tablet 14, the Sumerian uh (Akkadian kal-ma-tum) denotes the group containing insect
vermin. According to Landsberger’s (1934, 20-21) translation, the Sumerian uh.še, uh.še.kú and
uh.zí(d).da (Akkadian kal-mat še-im, ri-a-šu and kal-mat qé-mi) pertain to insect pests of grain and
flour, specifically barley. While Landsberger attempts to draw parallels between the uh.še.kú and
uh.zí(d).da and the Kornwurm and Melhwurm, respectfully (1934, 177), retranslation of the SumeroAkkadian text using Briggs et al. (2006) demonstrates that the transliteration is only capable of
referring to insect pests infesting stored grain (barely) and flour, making species level identification a
bit ambitious. The Sumerian determinative uh is often interpreted as louse. However, it is often used
in conjunction with the words for grain, fields, forest, etc. and, as such, should more accurately be
defined as vermin or pest (Tawil 1977). Uh combined with the word še, meaning barley or cereal,
denotes a pest of barley but does not specify the location/condition of the barley. The addition of the
kú, meaning fodder, in uh.še.kú versus its absence in uh.še in the Har-ra==Hubullu likely serves to
separate the pest of stored product barley, uh.še.kú, from the pest of field barley, uh.še. The position
of uh.še in the tablet following shortly after uh.a.šà(g).ga, rendered as a pest of the fields, further
supports its position as a pre-harvest pest rather than stored product. The tablet also lists a pest of
processed grain, i.e. flour—uh.zí(d).da, literally the insect pest with or in flour.
The Sumero-
Akkadians appear to have recognised three separate insect pests of cereals with preference to different
conditions of the cereal; pre-harvest, uh.še; post-harvest, unprocessed, uh.še.kú; and post-harvest,
processed, uh.zí(d).da. While the Har-ra==Hubullu provides insight into the habitat preference of
cereal pests, it does not provide the physical description necessary for identification of the insects.
The grain pests are also referred to in other literature of the period. For example, Brodenheimer
(1947) presents “A piece of linen is spread for a flea, a tissue for a moth, a granary for grain pests” as
a proverb with Sumero-Akkadian origins.
The earliest archaeological evidence [Table 5.1] for the presence of cereal pests in the region
can be dated to the Pre-Pottery Neolithic C period (PPNC), 8000-7500 BP. One of the earliest
accounts of grain pests comes from layer VI at Hacilar, SW Anatolia, dated to 7700-7550 BP (Helbaek
1970).
From structures Q.VI.1 and Q.VI.5, Helbaek describes the fragments of several adult
Sitophilus sp. in small heaps of charred wheat and barley. Additionally, one of the grains contained an
adult weevil and some kernels showed evidence of length-wise tunnelling.
As evidence of its
congeners S. oryzae and S. zeamais has yet to be recovered in the region at that date, the unidentified
Sitophilus sp. remains are likely to have been S. granarius. More conclusive evidence for the presence
of the granary weevil in the region at the time comes from a near contemporaneous well at Atlit-Yam
(circa 7500 BP) where Kislev et al. (2004) records 27 specimens of Sitophilus granarius.
118
Moving substantially forward in time, indirect evidence for the presence of S. granarius in
Assyrian and Hellenistic barley from Nimrud was referred to briefly by Helbaek (1970) as weevil
ravaged grain. Hopf and Zachariae (1921) found the grain weevil in 10th century BCE grain deposits
from Tel Arad in Northern Negev, and Kislev and Melamed (2000) reported insect remains, dating to
the 9th-10th century BCE, from charred grain and pulses found in store rooms near or in broken jars at
an Iron Age storage fort and village at Horbat Rosh Zayit, Israel.
Around 350 individuals of
Sitophilus granarius were recovered with or in wheat, Triticum parvicoccum, charred grain. Other
grain pests were noted: Alphitophagus bifasciatus, Oryzaephilus surinamensis (adult and pupa), and an
adult and whole larva of Tenebroides maritanicus.
5.3.2 Prehistoric Europe
The earliest evidence [Table 5.1] of cereal pests in Europe comes from a bandkeramic well in
Eythra village in Leipzig region of Germany. Well 2 (radiocarbon dated 7269-7180 BP) contained 204
individuals of Sitophilus granarius (Schmidt 2005). Schmidt has also recorded the granary weevil
from bandkeramic wells at Plaußig, dendrochonologically dated 7219 BP (2010a), from two wells at
Erkelenz-Kückhoven dated 7040 BP and 7007±5 BP (Schmidt 1998; 2010b), Eythra Well 1 dated
7034 BP (2005), and Köln approximately 6200 BP (1998). Büchner and Wolf (1997) have also found
Sitophilus granarius at Göttengen, Germany, 6030 BP.
While the granary weevil was clearly well established in northwestern Europe less than 500
years after its earliest fossil appearance to date in the Middle East, its path is unclear. It is obvious that
archaeological record is incomplete as the only other Neolithic account for Sitophilus granarius comes
from a cast in a piece of pottery from Servia (6700 BP), south Macedonia, Greece (Hubbard 1979).
While the archaeoentomological record for the granary weevil in Neolithic Europe is scarce, it is
practically non-existent for the other stored cereal pests. Valamoti and Buckland (1995) provide a
record of a Neolithic grain pest in Europe from Mandalo in western Macedonia, Greece, dating 5490 ±
55 BP (Kotsakis et al. 1989). A single charred head of Oryzaephilus surinamensis was recovered
from a large cache of emmer wheat during study of plant remains from the site. Additionally,
Tenebroides mauritanicus has been recovered from Erkelenz-Kückhoven (Schmidt 1998, 2010b) and
Plaußig (Schmidt 2010a) in Germany. T. mauritanicus has also been recorded from the German site
of Singen Offwiese (Schmidt 2007), which is associated with the Groß Gartach culture and dated 6950
BP (Dieckmann et al. 1997).
The paucity of archaeoentomological accounts for grain pests continues throughout the Bronze
and Iron Age. Fasani (1976) mentions the recovery of Sitophilus granarius from a Middle Bronze
Age site in Northern Italy, and the granary weevil was also present in Late Bronze Age France at the
site of Lake Bourget (Pecreaux 2008). Moreover, Rhyzopertha dominica has been recorded in Middle
Bronze Age contexts at Cova Punta Farisa in Fraga Huesca, Spain (Alonso and Buco 1993), and
119
Stegobium paniceum, has been found in Late Bronze Age Britain: Runnymede Bridge, Staines, Surrey
(Robinson 1991) and Wilsford, Wiltshire (Osborne 1989). The biscuit beetle has also been recovered
from an Iron Age site in Britain (Tattershall Thorpe, Lincolnshire, Chowne et al. 1986). Smith and
associates (2006) have found Sitophilus granarius in Iron Age contexts at Okruglo, Croatia, and
Compte and Perales (1984) have recovered the granary weevil, Rhyzopertha dominica, and Tribolium
sp. from Siriguarch, Alcañiz, Teruel, Spain.
5.3.3 Ancient Egypt
Like the Sumero-Akkadians, the ancient Egyptians left a scarce but usable documentary record
for the presence of grain pests. While Egyptian inscriptions have revealed images of invertebrates
which can be morphologically identified to genus (cf. Harpaz 1973; Levinson and Levinson 1998) the
ancient Egyptian language, like biblical Hebrew, lacked the generic term for ‘insect’. Depictions of
kheper beetles, scarabs, wasps, and locusts are clearly recognisable in glyphs and jewellery, but very
little distinction was made between worms, slugs, certain snakes, and holometabolic insect larvae. In
fact the Egyptian term h f a t (tola’ath in Hebrew) was indiscriminately used to denote all of the
aforementioned groups.
The Ebers Papyrus is an Egyptian medical document (c. 3552 BP) describing magical
formulas and remedies and is one of the earliest written records containing methods for deterring
pests. The Ebers Papyrus XCVIII provides instructions for controlling kkt-animals using burnt gazelle
dung diluted in water. Although Panagiotakopulu and others (1995) suggested that the use of kktanimals was a reference to grain weevils, it merely transliterates as ‘small animal’ and the species
identification is purely speculation based on context.
While species recognition through transliteration of hieroglyphs is tentative, the recovery of
palaeoentomological remains allows for more definite identification [Table 5.1].
Moreover, the
integration of archaeological material in addition to documentary evidence provides greater insight
into the presence and significance of grain pests in the region. The earliest archaeological records
come from Helbaek (cited in Solomon 1965), who notes Sitophilus granarius from the circa 4900 BP
Tomb in Saqqarah, and Solomon (1965), who mentions S. granarius having been recovered from the
tomb beneath the Step Pyramid of Saqqarah circa 4300 BP. The granary weevil and Stegobium
paniceum were recovered at the tomb of Queen Ichetis at Saqqarah, c. 4334-4150 BP (Chaddick and
Leek 1972).
A wheat deposit from a Middle Kingdom tomb at el-Gebelein (4181-4055 BP) yielded
Stegobium paniceum as well as the earliest fossil evidence for Trogoderma granarium
(Panagiotakopulu 2003).
Panagiotakopulu recognised five individuals of the khapra beetle and
seventeen individuals of the biscuit beetle. Tribolium sp. was recovered from a mid-3rd millennium
BC (5000-4000 BP) Egyptian tomb by Alfieri (in Andres 1931). Tribolium confusum was identified
120
from an offering pot from c. 3000 BP (Alfieri 1976), and Zacher (1937) recorded T. castaneum from
Egypt circa 3500 BP. Seifert (1987) reports the earliest record of Alphitobius diaperinus in an
unnamed New Kingdom Period site in Egypt.
Fossil evidence for Rhyzopertha dominica and Stegobium paniceum was available from
Liverpool Museum collections from Twelfth Dynasty Kahun, 3990-3800 BP (Panagiotakopulu 1998).
The R. dominica was recovered from a small sample of barley and is the earliest on record. The lesser
grain borer was also recovered in botanical remains from a vessel in Tutankhamun’s tomb, c. 3345 BP
(Alfieri 1931), and Zacher (1937) recorded T. castaneum, S. paniceum, Oryzaephilus surinamensis,
and Rhyzopertha dominica from another vessel from the tomb.
Material from the Workmen’s Village at Tell el-Amarna (thought to be dated between 33503323 BP based on pottery remains) contained grain pests from pigsty deposits. Panagiotakopulu
(1999) discusses the remains of S. granarius and Palorus ratzeburgi from coprolites at the site.
Additionally, Panagiotakopulu (2001) refers to Tribolium confusum, T. castaneum, Palorus
subdepressus and Cryptolestes turcicus as all having been recovered from Pharoanic Amarna. Zacher
(1934ab) notes Oryzaephilus surinamensis from a Minoan period vessel, 3350 BP.
Table 5.1 Archaeological sites presenting grain pests from Neolithic through Roman date contexts1
Location
Hacilar, SW
Anatolia
Atlit-Yam, Israel
Eythra village in
Leipzig region of
Germany; Well 2
Plaußig, Germany
ErkelenzKückhoven,
Germany, two
wells
Eythra, Germany
Well 1
Singen Offwiese,
Germany
Servia, south
Macedonia,
Greece
Köln, Germany
Göttengen,
Germany
Time, period
Pre-Pottery
Neolithic C,
7700-7550 BP
Pre-Pottery
Neolithic C,
circa 7500 BP
LBK, 7269-7180
BP
Species
Sitophilus granarius
Reference
Helbaek 1970
Sitophilus granarius
Kislev et al. 2004
Sitophilus granarius
Schmidt 2005
Sitophilus granarius,
Tenebroides
mauritanicus
Sitophilus granarius,
Tenebroides
mauritanicus
Schmidt 2010a
LBK, 7034 BP
Sitophilus granarius
Schmidt 2005
Neolithic, 6950
BP
Neolithic, 6700
BP
Tenebroides
mauritanicus
Sitophilus granarius
Dieckmann 1997;
Schmidt 2007
Hubbard 1979
LBK, 6200 BP
LBK, 6030 BP
Sitophilus granarius
Sitophilus granarius
Schmidt 1998
Büchner and
Wolf 1997
LBK, 7219 BP
LBK, 7040 BP
and 7007±5 BP
121
Schmidt 1998,
2010b
Mandalo in
western
Macedonia,
Greece
Tomb in
Saqqarah, Egypt
Egyptian tomb
Step Pyramid of
Saqqarah, Egypt
Tomb of Queen
Ichetis at
Saqqarah, Egypt
Tomb at elGebelein, Egypt
Kahun, Egypt
Egypt
Egypt
West House,
Akrotiri Santorini,
Thera
Knossos, Greece
Kommos, Greece
Northern Italy
Cova Punta Farisa
in Fraga Huesca,
Spain
Workmen’s
Village at Tell elAmarna, Egypt
Egypt
Tutankhamun’s
tomb, Egypt
Neolithic, 5490 ± Oryzaephilus
55 BP
surinamensis
Valamoti and
Buckland 1995
Early Dynastic
Period, circa
4900 BP
Early Dynastic
Period- Middle
Kingdom, 50004000 BP
Old Kingdom,
circa 4300 BP
Old Kingdom,
4334-4150 BP
Sitophilus granarius
Helbaek cited in
Solomon 1965
Tribolium sp
Alfieri cited in
Andres 1931
Sitophilus granarius
Solomon 1965
Sitophilus granarius,
Stegobium paniceum
Chaddick and
Leek 1972
Stegobium paniceum,
Trogoderma
granarium
Rhyzopertha
dominica, Stegobium
paniceum
Panagiotakopulu
2003
Alphitobius diaperinus
Seifert 1987
Tribolium castaneum
Zacher 1937
Sitophilus granarius,
Rhyzopertha
dominica, Stegobium
paniceum,
Oryzaephilus sp.
Sitophilus granarius
Panagiotakopulu
and Buckland
1991
Jones 1984
Sitophilus granarius,
Tribolium confusum
Sitophilus granarius
Shaw and Shaw
1995
Fasani 1976
Rhyzopertha dominica
Alonso and Buxo
1993
Sitophilus granarius,
Palorus ratzeburgi
Panagiotakopulu
1999
Oryzaephilus
surinamensis
Rhyzopertha
dominica, Tribolium
castaneum, Stegobium
paniceum,
Oryzaephilus
surinamensis
122
Zacher 1934ab
Middle
Kingdom, 41814055 BP
Middle
Kingdom,
Twelfth Dynasty,
3990-3800 BP
New Kingdom,
3520- 3020 BP
New Kingdom,
circa 3500 BP
Late Minoan,
circa 3500 BP
Late Minoan, c.
3425 BP
Late Minoan, c.
3425 BP
Middle Bronze
Age, 3450-3250
BP
Middle Bronze
Age, 3450-3250
BP
New Kingdom,
3350-3323 BP
New Kingdom,
3350 BP
New Kingdom,
c. 3345 BP
Panagiotakopulu
1998
Alfieri 1931;
Zacher 1937
Wilsford,
Wiltshire, UK
Late Bronze
Stegobium paniceum
Age, 3330 ± 90
BP
Lake Bourget,
Late Bronze
Sitophilus granarius
Savoie, France
Age, 3250-2750
BP (Gauthier and
Richard 2009)
Amarna, Egypt
Pharaonic
Tribolium confusum,
T. castaneum, Palorus
subdepressus,
Cryptolestes turcicus
Egypt
Third
Tribolium confusum
Intermediate,
Twenty-first
Dynasty, 3000
BP
Nimrud
Assyrian and
Sitophilus granarius
Hellenstic
Periods, 39002273 BP
Runnymede
Late Bronze
Stegobium paniceum
Bridge, Staines,
Age, 2950-2800
Surrey, UK
BP
Tel Arad in
Iron Age, 2950Sitophilus granarius
Northern Negev
2851 BP
Horbat Rosh
Iron Age, 2950Sitophilus granarius,
Zayit, Israel
2751 BP
Alphitophagus
bifasciatus,
Oryzaephilus
surinamensis,
Tenebroides
mauritanicus.
Okruglo, Croatia
Iron Age, 2760Sitophilus granarius
2640 BP
Laemophloeus
(=?Cryptolestes) spp.
Siriguarach,
Iron Age, 2650Sitophilus granarius,
Alcañiz, Teruel,
2551 BP
Rhyzopertha
Spain
dominica, Tribolium
sp.
Tomb from Hunan 2100 BP
Trogoderma persicum,
Province in China
Sitophilus oryzae
Tattershall
Iron Age, 2350 ± Stegobium paniceum
Thorpe,
90 BP
Lincolnshire, UK
Santa Pola, Spain Roman, 1950Sitophilus granarius
1851 BP
(1st c. AD)
Alphen aan den
Roman, 1950Sitophilus granarius
Rijn, Netherlands 1851 BP
(1st c. AD)
Neuss
Roman, 1950Sitophilus oryzae
(Novaesium IV),
1900 BP
Germany
(early 1st c. AD)
123
Osborne 1989
Pecreaux 2008
Panagiotakopulu
2001
Alfieri 1976
Helbaek 1970
Robinson 1991
Hopf and
Zachariae 1921
Kislev and
Melamed 2000
Smith et al. 2006
Compte and
Perales 1984
Chu and Wang
1975
Chowne et al.
1986
Moret and Martin
Cantarino 1996
Kuijper and
Turner 1992
Knörzer 1970
Neuss
(Novaesium),
Germany
1 Poultry, Central
London, UK
Roman, 1920 BP
(30 AD)
Roman, 19031890 BP
(47-60 AD)
21 Saint Peters
Street, Colchester,
UK
Roman, 18881870 BP
(62-80 AD)
Coney Street,
York, UK
Roman, 18791876 BP
(71-74 AD)
Touffréville
Calvados, France
Roman, circa
1875 BP
(c. 75 AD)
Herculaneum,
Naples, Italy
Roman, circa
1871 BP
(c. 79 AD)
Roman,
approximately
1875- 1800 BP
(late 1st to mid2nd c. AD)
Papcastle,
Cumbria, UK
Amiens, France
Roman, c. 18491750 BP
(2nd c. AD)
Mons Claudianus,
Egypt
Roman, c. 18491750 BP
(2nd c. AD)
Roman, c. 17751750 BP
(late 2nd c. AD)
Woerden, ZuidHolland
Oryzaephilus
surinamensis,
Sitophilus granarius
Sitophilus granarius,
Oryzaephilus
surinamensis,
Cryptolestes
ferrugineus, Palorus
ratzeburgi
Sitophilus granarius,
Oryzaephilus
surinamensis,
Cryptolestes
ferrugineus, Palorus
ratzeburgi
Tenebroides
mauritanicus,
Cryptolestes
ferrugineus,
Oryzaephilus
surinamensis, Palorus
ratzeburgi, Tenebrio
obscurus, Sitophilus
granarius
S. granarius,
Stegobium paniceum,
Tenebrio obscurus,
Oryzaephilus sp.
Sitophilus granarius,
Oryzaephilus sp.
Cymorek and
Koch 1969; Koch
1970
Rowsome 2000;
Smith in press
Cryptolestes
ferrugineus,
Oryzaephilus
surinamensis, Palorus
ratzeburgi,
Alphitobius
diaperinus, Sitophilus
granarius
Stegobium paniceum,
Cryptolestes
ferrugineus,
Oryzaephilus
surinamensis,
Tenebrio sp., Palorus
ratzeburgi, Sitophilus
granarius
Oryzaephilus sp. and
Cryptolestes turcicus
Kenward and
Allison 1995
Sitophilus granarius,
Oryzaephilus
surinamensis,
124
King and Hall
2008
Hall and
Kenward 1976;
Kenward and
Williams 1979
Ponel et al. 2000
Dal Monte 1956
Matterne et al.
1998; Yvinec
1997
Panagiotakopulu
and van der Veen
1997
Pals and Hakbijl
1992
Cryptolestes
ferrugineus, Palorus
ratzeburgi, Tenebrio
molitor,
Alphitophagus
bifasciatus
Skeldergate Well, Roman, 1750 ±
Sitophilus granarius,
York, UK
80 BP
Cryptolestes
ferrugineus, Palorus
(c. 200 AD)
ratzeburgi, Tribolium
castaneum, Tenebrio
obscurus,
Oryzaephilus
surinamensis,
Stegobium paniceum
Alcester,
Roman, 1750Sitophilus granarius,
Warwickshire,
1651 BP
Palorus subdepressus,
UK
(3rd c. AD)
Tenebrio obscurus,
Oryzaephilus
surinamensis,
Stegobium paniceum
Hambacher Forest Roman, 1712 ± 5 Sitophilus granarius
near Köln,
BP
Germany
(c. 238 AD)
Towcester,
Roman, 1620Oryzaephilus
Northamptonshire, 1585 BP
surinamensis,
UK
(330-365 AD)
Cryptolestes turcicus,
Cryptolestes
ferrugineus, Tribolium
castaneum, Tribolium
confusum, Sitophilus
granarius
1
Hall et al. 1980
Osborne 1971
Schmidt 2006b
Girling 1983
Table does not contain a comprehensive list of Romano-British sites containing grain pests
5.3.4 Ancient Greece and Aegean
The Greek names κίς, κορίς, φθείρ, σής, ïψ, σκυίψ and θρίψ refer to small insect pests
(Beavis 1988). Κίς, in particular, is normally restricted to insects infesting grains or pulses. Although
Κίς is often read as weevil by modern scholars, inferring Sitophilus granarius, the term, like the
Sumero-Akkadian uh.še.kú, comprised a number of modern grain species, and in Suidas’ lexicon is
simply defined as vermis, or vermin. The Greeks, like the earlier writers, do not offer a description of
the insect pests, and as such, the terms could refer to any number of stored product insects. For
example, Theophrastus (2321-2236 BP) CP IV.15.6 has been translated as “Each kind of seed-crop
when put under a roof produces from its proper fluidity certain animals of a form peculiar to itself: so
wheat and barley produce their weevils…” (1990, 355); whereas it may be more accurate if the term
‘weevils’ was replaced by ‘grain pests’.
125
The fossil record for pre-Roman Greece and Aegean Islands is remarkably slight [Table 5.1].
A carbonised Sitophilus granarius was found in a sample of barley from a destruction deposit at the
‘unexplored’ mansion complex at Knossos, with a Late Minoan date, around 3425 BP (Jones 1984),
and Shaw and Shaw (1995) report S. granarius and Tribolium confusum from a contemporaneous site
at Kommos. The remains of Sitophilus granarius, Rhyzopertha dominica, Stegobium paniceum, and
Oryzaephilus sp. were identified in samples from the West House, Akrotiri Santorini, Thera circa
3500 BP (Panagiotakopulu and Buckland 1991).
5.3.5 China
The palaeoentomological record for the Orient is very limited [Table 5.1]. In regards to stored
product pests, Chu and Wang (1975) reported Trogoderma persicum (Pic.) (i.e. T. variabile Ball.) and
Sitophilus oryzae from a tomb dated about 2100 BP in Hunan Province in China. Archaeological
excavation accompanied by bioarchaeological analysis will surely produce many ancient records of
pests and other insects in Asia and the Indian subcontinent in due course.
5.3.6 The Roman Period
In comparison to earlier periods, the Roman authors wrote prolifically about grain pests
starting with Marcus Porcius Cato’s 2185 BP De Re Rustica (Ag. XCII), and seemed particularly
concerned with a grain pest known as curculio. In his poem, Virgil exclaims “populatque ingentem
farris acervum curculio” (G. I.CLXXXVI)—a huge grain (spelt)-heap curculio ravages, and Plautus’
play Curculio features a character named Curculio Parastus (parasite), who is portrayed as a greedy,
gluttonous, and unscrupulous character. In his opening lines, Curculio utters:
“Make way for me, friends, strangers, while I do my duty here! Scatter, clear out, get
off the street, everybody, so that I may not career into anyone and lay him out with
my head, or elbow, or chest, or knee! I tell you what, it’s a sudden, pressing, urgent
job I’m charged with now, and there’s no man rich enough to block my path—neither
general, nor despot, any of ‘em, nor market inspector, no mayor, nor burgomaster, I
don’t care how grand he is—down he’ll go, down he’ll drop from the sidewalk and
stand on his head in the street!” (PC, 219-221).
It appears that curculio was perceived as an insect that was capable of ravaging enormous piles of
grain and infesting the cereals of people regardless of societal position.
Moreover, the Roman writers seem to differentiate between curculio and other grain-associated
insects, as indicated in Vitruvius’s commentary on architecture “granaria sublimata et ad
septentrionem aut aquilonem spectantia disponantur, ita enim frumenta non poterunt cito concalescere,
sed ab flatu refrigerata diu servantur. namque ceterae regiones procreant curculionem et reliquas
bestiolas quae frumentis solent nocere,” (Arch VI.VI.IV)—the granaries are raised, and must be
126
towards the north or east, so that the grain may not heat, but be preserved by the coolness of the air; if
towards other aspects, curculio, and other insects that harm grain, will be generated. Although the
Romans recognized different species of grain pests, only curculio was named, and as in previous
periods, the species, unfortunately, lacks a physical description making species level identification
difficult.
The majority of the references to curculio concern agriculture or natural history, in which the
authors discuss methods for preventing the contamination of cereals and purging an infestation. A
brief investigation of a few these works is worthwhile as the authors provide insight into the behaviour
of curculio as well as a few locations in which it was found.
Approximately 1986 BP, Varro (as
translated by Hooper and Ash 1935) writes:
"Wheat should be stored in granaries, above ground, open to the draught on the east
and north, and not exposed to damp air rising in the vicinity. The walls and floor are
to be coated with marble cement, or at least with clay mixed with grain-chaff and
amurca, as this both keeps out mice and worms and makes the grain more solid and
firm. Some farmers sprinkle the wheat, too, with amurca, using a quadrantal to about
a thousand modii.
Different farmers use different powders or sprays, such as
Chalcidian or Carian chalk, or wormwood, and other things of this kind. Some use
underground caves as granaries, the so called sirus, such as occur in Cappadocia and
Thrace; and still others use wells, as in the Carthaginian and Oscensian districts in
Hither Spain. They cover the bottom of these with straw, and are careful not to let
moisture or air touch them, except when the grain is removed for use; for [curculio]
does not breed where air does not reach. Wheat stored in this way keeps as long as
fifty years, and millet more than a hundred. Some people, as in Hither Spain and in
Apulia, build granaries in the field, above ground, so constructed that the wind can
cool them not only from the sides, through windows, but also beneath from the
ground. […] Grain which [curculio] has begun to infest should be brought out for
protection. When it is brought out, bowls of water should be placed around in the
sun; [curculio] will congregate at these and drown themselves. Those who keep their
grain under ground in the pits which they call sirus should remove the grain some
time after the pits are opened, as it is dangerous to enter them immediately, some
people having been suffocated while doing so. Spelt which you have stored in the ear
at harvest-time and wish to prepare for food should be brought out in winter, so that it
may be ground in the mill and parched.” (RR I.LVII 1935).
Around 1900 BP, Columella (as translated by Ash1941) writes:
“And I am not unaware that some consider the best place for storing grain to be a
granary with a vaulted ceiling, its earthen floor, before it is covered over, dug up and
127
soaked with fresh and unsalted lees of oil and packed down with rammers as is
Signian work. Then, after this has dried thoroughly, it is overlaid in the same way
with a pavement of tiles consisting of lime and sand mixed with oil lees instead of
water, and these are beaten down with great force by rammers and are smoothed off;
and all joints of walls and floor are bound together with a bolstering of tile, for
usually when buildings develop cracks in such places they afford holes and hidingplaces for underground animals. But granaries are also divided into bins to permit
the storage of every kind of legume by itself. The walls are coated with a plastering
of clay and oil lees, to which are added, in place of chaff, the dried leaves of the wild
olive or, if these are wanting, of the olive. Then, when the aforesaid plastering has
dried, it is again sprinkled over with oil lees: and when this has dried the grain is
brought in. This seems to be the most advantageous method of protecting stored
produce from damage by [curculio] and like vermin, and if it is not carefully laid
away they quickly destroy it. But the type of granary just described, unless it be in a
dry section of the steading, causes even the hardest grain to spoil with mustiness; and
if it were not for this, it would be possible to keep grain even buried underground, as
in certain districts across the sea where the earth, dug out in the manner of pits, which
they call siri, takes back to itself the fruits which it has produced. But we, living in
regions which abound in moisture, approve rather the granary that stands on supports
above the ground and the attention to pavements and walls as just mentioned,
because, as I have said, the floors and sides of storerooms so protected keep out
[curculio]. Many think that when this kind of pest appears it can be checked if the
damaged grain is winnowed in the bin and cooled off, as it were. But this is a most
mistaken notion; for the insects are not driven off by so doing, but are mixed through
the whole mass. If left undisturbed, only the upper surface would be attacked, as
[curculio] breeds no more than a palm's breadth below; and it is far better to
endanger only the part already infested than to subject the whole amount to risk. For
it is easy, when occasion demands it, to remove the damaged portion and use the
sound grain underneath. But these latter remarks, though brought in extraneously, I
nevertheless seem to have introduced not unseasonably at this point.” (RR I.VI.XVXII 1941).
The commentaries of Varro and Columella seem to depict curculio as a primary pest by
insinuating that the species was responsible for infesting undamaged grains.
Based on
archaeoentomological accounts, the Mediterranean region was infested by two primary pests of grains,
Sitophilus granarius and Rhyzopertha dominica. As R. dominica does not appear to have had a
widespread distribution in the past (though this may be a reflection of a poor fossil record), the Roman
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authors may have been referring to Sitophilus granarius, which is the species most commonly
associated with the term curculio by modern scholars in translating it as weevil. However, the
possibility cannot be completely discounted of curculio having been used in reference to both species.
Varro specifies several regions—Cappadocia, Thrace, Carthaginian and Oscensian districts in
Hither Spain, and Apulia—that sought to establish a means of pest control against infestations of
curculio. As there is not presently an established fossil record for 1986 BP, Varro’s De Re Rustica
provides the best evidence for the distribution of curculio at that time. Strabo’s description of
Cappadocia places the region in modern-day central Anatolia where it extended northwards to the
Black Sea from Mount Taurus, east to the Euphrates and west to Lake Tuz. Thrace refers to the region
north of Thessaly extending northwards to the Hæmus Mons (Balkan Mountains), west to the Nestos
(Mesta) River, and east to the Black Sea (Carr 1838). Hither Spain, most likely, refers to Hispania
Citerior, which included the northeastern coast and Ebro valley of modern Spain, and Strabo’s
description of Apulia places the area in southeastern Italy (in the heel-region of the boot), bordering
the Adriatic and Ionian Seas (SG VI.III). Varro’s reference implies that curculio was established
along most of the Mediterranean coast of southern Europe during the Roman Republic.
First Century AD
Records of insect assemblages from Continental Europe in the Roman period are rare and there
is a dearth of observations of grain pests [Table 5.1]. The earliest remains of Roman age grain pests
are dated to 30 AD, Oryzaephilus surinamensis and Sitophilus granarius from Neuss (Novaesium) in
Germany (Cymorek and Koch 1969; Koch 1970), and early first century AD contexts from NeusNovaesium IV, Germany have provided Sitophilus oryzae in association with charred rice (Knörzer
1970). In France, Ponel and associates (2000) noted S. granarius, Stegobium paniceum, Tenebrio
obscurus, and Oryzaephilus sp. from Touffréville Calvados, dated circa 75 AD, and Dal Monte (1956)
recorded larval, pupal and adult Sitophilus granarius as well as a single Oryzaephilus sp. in infested
charred wheat from beneath the AD 79 tephra at Herculaneum, Naples, Italy. S. granarius was also
recovered from contexts at Alphen aan den Rijn, Netherlands dating to the first century AD (Kuijper
and Turner 1992) as well as Valkenburg fort (Hakbijl 1988).
The archaeoentomological evidence suggests that the grain pests entered Britain immediately
following the arrival of the Roman legions; there are no records to indicate the presence of grain pests
in Britain prior to the arrival of the Roman military forces. Buildings, workshops, yards and pits at
Poultry, Central London, were dated from just after 47 AD, the start of the occupation of Roman
London (Rowsome 2000; Smith in press), and the deposits were all sealed by the 60 AD ‘fire horizon’
interpreted as the burning of London during the Boudiccan revolt (Rowsome 2000). From the Poultry
site samples, Smith (loc. cit.) identified myriad insect remains including Sitophilus granarius,
Oryzaephilus surinamensis, Cryptolestes ferrugineus, and Palorus ratzeburgi. The same range of
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species was present in the timber drain at 21 Saint Peters Street, Colchester, constructed immediately
following the Boudiccan revolt (King and Hall 2008).
In northern England, grain pests have been noted in the Roman Fort at the Millennium site at
Carlisle Castle 72/3 AD (Smith unpublished), at other sites in and near the Roman fortress at Carlisle
(Kenward et al. 2000; Kenward and Carrott 2006), and the fort at Ribchester, Lancashire 71-4 AD
(Large et al. 1994; Buxton and Howard-Davis 2000). The humic silts from in and around the beam
slots of a late first century wooden building at Coney Street, York produced immense numbers of
grain pests associated with spoilt grain. The humic silt layer 2105 yielded a range of grain fauna—
Tenebroides mauretanicus, Cryptolestes ferrigineus, Oryzaephilus surinamensis, Palorus ratzeburgi,
Tenebrio obscurus, and Sitophilus granarius—representing a massive infestation (Hall and Kenward
1976; Kenward and Williams 1979).
Second, Third, and Fourth Centuries AD
While there is a dearth of information concerning the Italian sites, the grain pests continue to
be represented in France, Germany, and England beyond the first century [Table 5.1]. In France, a
Gallo-Roman granary burned during the second century in Amiens provided a fauna which included
Stegobium paniceum, Cryptolestes ferrugineus, Oryzaephilus surinamensis, Tenebrio sp., Palorus
ratzeburgi, and Sitophilus granarius (Yvinec 1997; Matterne et al. 1998), and Schmidt (2006)
demonstrates the presence of S. granarius in the second century of Hambacher Forest near Köln in
Germany.
In Britain, grain associated insect faunas have been recorded from numerous sites
throughout England (e.g. Buckland 1982; Kenward and Carrott 2006) and as far north as Invereskgate
in West Lothian, Scotland (Smith 2001) until the end of the 4th century AD. The second century
provides evidence for the arrival of Alphitobius diaperinus (Kenward et al. 1986; Hall and Kenward
1990; Kenward and Allison 1995; Kenward et al. 2000), Stegobium paniceum (Hall and Kenward
1990), and Tribolium castaneum (Hall et al. 1980; Hall and Kenward 1990) in Northern England.
Moreover, the third and fourth centuries AD indicate the introduction of Palorus subdepressus
(Osborne 1971) as well as Cryptolestes turcicus and Tribolium confusum (Girling 1983), respectively,
to England. On the other side of the Mediterranean, Oryzaephilus sp. and Cryptolestes turcicus have
been recovered from a second century quarry site at Mons Claudianus in Egypt (Panagiotakopulu and
van der Veen 1997).
Additionally, the Roman period provides the earliest archaeological evidence for the
movement of grain pests via ships. Pals and Hakbijl (1992) report Sitophilus granarius, Oryzaephilus
surinamensis, Cryptolestes ferrugineus, Palorus ratzeburgi, Tenebrio molitor, Alphitophagus
bifasciatus, and the parastoid wasp Lariophagus distinguendus (Förster) from the remains of a late
second century ship near the presumed Roman fort of Laurium in Woerden, Zuid-Holland. Similarly,
Rule and Monaghan (1993) have reported Sitophilus sp. (likely S. granarius) from the third century
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ship wreck near Guernsey. At minimum, the present of pests in these vessels reveals that grain was
still a traded commodity in the Roman Empire during the second and third centuries.
5.4 Discussion: The Origins and Diffusion of the Species
While a few early writers acknowledged the depredations of the various grain fauna dating
back to the Hammurabian period, the literature does not allow conclusive reconstruction of species’
distributions, as pointed out by Buckland (1981), even when species level identification is discernable
from the texts. Thus the early literary accounts are not viable alternatives to archaeological and
palaeoecological data when assessing dispersal patterns of the cereal pests. In this section, the
diffusion of the grain-associated entomofauna will be assessed through an evaluation of spatial and
temporal changes in the various species, i.e. biogeography.
5.4.1 Neolithic
At present, the earliest archaeological evidence of farming communities and founder crops
associated with Neolithic agriculture dates to c. 12000 BP in southwestern Asia (Colledge et al. 2004),
c. 8350 BP in southern Greece (Perlès 2001), and c. 7450 BP in Germany (Stäuble 1995). As early as
the 7th millennium BP, the archaeoentomological evidence, predominantly from Sitophilus granarius,
indicates the presence of pests in three regions: Germany, Macedonia, and the Middle East. In the
Middle East, there is approximately a 4,000 year gap between the earliest archaeological evidence for
the emergence of farming communities and the first indication of grain-associated insects. Moreover,
while a difference of circa 1700 years exists in the Greco-Macedonian region, the grain pests are
seemingly present almost immediately (c. 7269 BP) following the agriculturalisation of Germany. Did
the grain pests make the transition to synanthropy independently in separate locations or in a single
region and disperse via anthropic transportation?
Currently, the earliest fossil record of grain pests, i.e. S. granarius, dates to 7700-7550 BP in
southwest Anatolia (Helbaek 1970). As this early record pre-dates the emergence of agriculture in
northwest Europe by approximately 300 years and the earliest German specimens by circa 500 years,
it likely signifies the region of origin in which the species became associated with human-stored
cereals. There has been an unfortunate paucity of archaeoentomological investigations undertaken in
the Middle East, particularly in regards to the early Neolithic, and even the earliest grain pest
accounts, i.e. Helbaek 1970 and Kislev et al. 2004, were stumbled upon haphazardly during
archaeobotanically aimed investigations of charred material.
Whereas the majority of
archaeobotanical studies may have overlooked or not been concerned with the recovery of insect
remains, even the few archaeoentomologically aimed assessments, e.g. G. A. King’s (unpublished)
sampling of Pre-Pottery Neolithic A and B contexts at Catalhöyük, have not recovered evidence of
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grain fauna prior to the Pre-Pottery Neolithic C (PPNC). Despite this, it would be premature to
completely dismiss an earlier Middle Eastern presence as future investigations may reveal earlier
evidence for grain pests in the region.
The initial introduction of the grain fauna and their subsequent dispersal in the Middle East
would have been inhibited by factors during the early Neolithic. The granary weevil is the sole fossil
evidence for cereal pests in the Pre-Pottery Neolithic C Middle East, and, as stated above, the species
is flightless and regarded as a poor disperser (Mlambo 1980). Because of its hypothesised commensal
relationship with pre-Neolithic rodents, S. granarius would have likely been introduced into the manmade grain stores via their associated rodents. When the rodents began constructing their dens in
anthropic environments (zooarchaeological accounts suggest this move towards commensalism occurs
as early as the PPNA (11500-10500 BP) with the house mouse, Mus musculus, Bar-Yosef et al. 1991;
Tchernov 1994; Jenkins 2003), the granary weevils would have been able to make the transition from
the rodent stores to the similar microhabitat found in the grain stores of man. This would suggest that
the S. granarius may be present in archaeological contexts dating to the PPNA.
Additionally,
excavations at Dhra’, near the Dead Sea in Jordan, have revealed evidence of 3 X 3m circular
structures interpreted as extramural (located between buildings) granaries, dated to 11300-11200 BP
(Kuijt and Finlayson 2009). These granaries suggest that cereals may have initially been owned
communally, but by 10500 BP, there is a shift to small-scale, house-hold level storage (Kuijt and
Finlayson 2009). If the grain fauna were present in the region during the early PPNA, there may be
evidence of infestations in the extramural granaries; however, the later adoption of small-scale storage
of grain would have reduced the chances of pest infestation and the archaeological presence of the
grain pests. Moreover, dispersed settlement patterns and lack of settlement continuity would have
limited or contained the diffusion of early pest infestations (Buckland 1990).
Alternatively, ecological constraints may not have presented the opportunity for the initial
introduction of the granary weevil until the Late PPNB- Early PPNC.
At this date, the large
agricultural villages along the Mediterranean zone of the Jordan Valley shifted to the eastern side of
the valley into Mediterranean and desert areas; many of these new settlements are larger than previous
villages and are founded in locations with no evidence of earlier PPNB occupation (Rowan and
Golden 2009). Following this shift, there is an apparent contraction in population (Rowan and Golden
2009). Around 8200 BP, paleoclimate studies (e.g. Berger and Guilaine 2009; Magney et al. 2003;
Figure 5.1) indicate a drastic change in the region’s climate towards a more temperate zone in
Anatolia and northern Israel, which is favoured by the granary weevil (Kislev et al. 2004). If the grain
pests had not made the transition to man-made grain stores by the mid-PPNB, the combination of
elements present during the Late PPNB-PPNC would have favoured its infestation.
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Figure 5.1 Climate zones across Eurasia and North Africa around 8200 BP (Berger and Guilaine 2009; Magney et al.
2003); the green circles represent arid climate and fresh for northern Europa; yellow circles indicate a very contrasted
climate; and blue circles denote a wet and fresh climate
Unlike the Middle East, there have been a number of archaeoentomological investigations
carried out on Neolithic, early Linearbandkeramik (LBK), sites in northwestern Europe, particularly
Germany. These investigations have demonstrated the presence of grain pests in some of the earliest
sites with evidence of neolithization. The recovery of Sitophilus granarius from sites dating to 7269
BP at Eythra (Schmidt 2005) in Neolithic Germany comprehends a demic expansion of agriculture,
i.e. through the immigration of people (cf. Childe 1925), resulting from the anthropic transportation of
pest infested cereals during population movement or grain trade rather than through cultural
transmission, i.e. the adoption of cultural traits not necessarily associated with the long-distance
movement of individuals (cf. Whittle 1996), and the multi-regional synanthropic transition of the
granary weevil. The hypothesis of Sitophilus granarius as an introduced species is supported by
archaeological, ecological and geographical factors:
1) The myriad ecological and geographical barriers between the Middle East and northwestern
Continental Europe, e.g. the Black Sea, the Carpathian Mountains, and the Mátra Mountains,
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would not have provided a corridor pathway by which the flightless weevil could have freely
dispersed and populated the European regions naturally;
2) The Near East fossils are morphologically similar to the remains recovered in the German
contexts.
If the Sitophilus populations had evolved independently through geographic
isolation, vicariance would be expected; as theoretically the subdivision of the geographic area
would have contributed to the splitting of the taxon. However, genetic analysis should be
adopted to explore this possibility;
3) According to the present zooarchaeological record, commensal rodents do not arrive in the
region until 6450 BP (Auffray et al. 1990; Sommerville 1999) with the recovery Mus musculus
from Place St-Lambert, Liège Belgium (Cordy and Stassart 1982). Without the presence of
commensal rodents to provide a stepping stone towards synanthropy, the weevils would have
had to have been introduced via another route; in the absence of indigenous cereal progenitor
species, perhaps through the initial collection and storage of infested acorns or seeds.
However, a population adapted to the long-term infestation of acorns or seeds would be
morphologically larger than a population adapted to small cereal grains.
In addition to the recovery of Sitophilus granarius, another grain pest, Tenebroides
mauritanicus, has been recovered in the region as early as 7219 BP at Plaußig (Schmidt 2010a) and
slightly later at Erkelenz-Kückhoven (Schmidt 1998, 2010b). While T. mauritanicus may have been
imported into Germany alongside the granary weevil, the absence of earlier fossil evidence supports
the idea that it is likely endemic to Europe. Unlike Sitophilus granarius, which develops entirely
within the confines of the cereal kernels and thus can be unwittingly transported by man, the cadelle is
one of the largest stored product beetles. As an adult it can grow to a length of 11 mm, and its larvae
can reach up to 20 mm (Mound 1989). During the small-scale transportation of cereals, it would have
been a visible contaminant (though perhaps not a concern to Neolithic people). Although largely
synanthropic today, Palm (1959) has recorded the cadelle under beech bark in southern Europe. The
transition from under bark pabulum to grain storage has been discussed above for Oryzaephilus,
Cryptolestes, and Palorus. Tenebroides mauritanicus, as a scavenger and predator, would have been
well adapted for the micro-habitat of the grain storage environment.
Alternatively, as the species has been recovered from Iron Age sites in the Near East, e.g.
Horbat Rosh Zayit, Israel (Kislev and Melamed 2000), it may have been indigenous to that region and
dispersed alongside Sitophilus granarius. The temperature requirements required for the completion
of the cadelle’s life cycle closely resemble that of Sitophilus granarius [Figure 5.2], with both species
requiring temperatures above 15 °C. Palaeoecological studies on the 8200 BP climate event suggest
that the summertime temperatures in northwestern Europe were only around 1 °C (Berger and
Guilaine 2009), which would not have met the needs of the species prior to their synanthropic
transition. Additionally, Klitgaard-Kristensen et al. (1998) observed a reduction in the tree-ring
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growth of oaks from Bamberg Germany circa 8200 BP, which may be attributed to lower summer
time temperatures.
Despite the absence of fossil evidence, the cadelle’s temperature requirements
suggest that the climate in Germany around 8200 BP would not have been conducive to supporting
viable populations of the cadelle. As such, it was most likely anthropically transported into the region
from the Middle East alongside the granary weevil. However, the tree-ring studies by KlitgaardKristensen et al. (1998) as well as oxygen-isotope ratios from deep-sea ostracods from the Amersee in
southern Germany (von Grafenstein et al. 1998) propose that the low temperatures may have been the
result of rapid climatic change. In the absence of a fossil record, the possibility that T. mauritanicus
occupied the region prior to the decrease in temperatures, survived in refugia, and reoccupied
afterwards must be considered.
The MCR data calculated using insect remains recovered
archaeologically from well contexts in Germany, which were contemporaneous to the cadelle’s
Neolithic presence, support the possibility of non-synanthropic populations by 7200 BP. The MCRs
indicate that the temperatures during the warmest month were above 15 °C, which falls within the
thermal range for the species to complete its life cycle [Appendix 1D]. The lack of fossil record could
be attributed to 1) the few archaeoentomological surveys conducted on Neolithic sites in the Middle
East and 2) the large size of the beetle resulting in the increased fragmentation of remains and
problematic preservation (Kenward pers. comm).
The Neolithic presence of grain fauna in Macedonia is late in comparison to the arrival of
agriculture in the region. Moreover, the evidence is meager, consisting of an imprint of Sitophilus
granarius from a piece of pottery in Servia (Hubbard 1979) and a single charred head of Oryzaephilus
surinamensis from an emmer cache in Mandalo (Valamoti and Buckland 1995). If we assume that the
cast represents the presence of granary weevil populations in the region, the Greek Sitophilus
granarius could present a similar scenario to the German specimens and infer the importation of
cereals into the area from the Near East. Otherwise the populations would have been geographically
separated by mountains and waterways, which would have restricted the diffusion of the species
naturally, and vicariance would be present. However, because of the paucity of archaeoentomological
and contextual evidence, it would be ill-advised to discount the possibility that the granary weevil was
native to the region.
Alternatively, as the evidence is based solely on a cast, the presence of the granary weevil in
Macedonia cannot be irrefutably confirmed as the pottery may even have been imported into the
region. The absence of S. granarius in Greece would be significant as it could indicate that agriculture
was initially introduced into the region through cultural transmission. This would explain the absence
of S. granarius from the large emmer cache at Mandalo, which contained the saw-toothed grain beetle;
a concern noted by Valamoti and Buckland (1995). The Oryzaephilus specimen most likely represents
an autochthonous component, introduced from the local environment as a mould-feeder.
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The archaeoentomological evidence for the Neolithic is limited. However, a biogeographical
interpretation of the remains permits speculation in regards to the spread of agriculture. The recovery
of the granary weevil from the Near East and northwest Europe supports the argument for a demic
expansion of agriculture with a direct connection between the regions; whereas the Greek material
could infer the introduction of agriculture from the Near East largely through cultural transmission,
but the evidence is fairly inconclusive.
5.4.2 The Bronze Age
While the Neolithic heralded the emergence and diffusion of agriculture, the Bronze Age,
noted archaeologically for metalworking involving the use of bronze alloy, witnessed the rise of tradenetworks. Archaeoentomologically, the Bronze Age provides evidence towards the emergence of
grain-associated faunas initially in Egypt and later in the Aegean and southern Europe. Although
archaeological evidence shows that crops had been domesticated ubiquitously around the
Mediterranean by this date, the archaeoentomological evidence suggests that cereals were being
transported, if not exchanged, during the Bronze Age.
G r a in A sso c ia t e d S p e c ie s
Figure 5.2 Range (light) and optimal (dark cross-hatching) ranges of temperature °C, which are required for the
completion of developmental life cycle, for grain fauna
Alphitobius diaperinus
Alphitophagus bifasciatus
Cryptolestes ferrugineus
Cryptolestes turcicus
Oryzaephilus surinamensis
Palorus ratzeburgi
Palorus subdepressus
Rhyzopertha dominica
Stegobium paniceum
Sitophilus granarius
Sitophilus oryzae
Sitophilus zeamais
Tenebrio molitor
Tenebrio obscurus
Tenebroides mauritanicus
Tribolium castaneum
Tribolium confusum
Trogoderma granarium
Trogoderma persicum
0
5
10
15
20
25
30
35
40
45
50
Temperature °C
Sources consulted: Sinha and Watters 1985; Sinha 1991; Semple et al. 1992; Rees 2007
The earliest Bronze Age account of grain pests, i.e. Sitophilus granarius, dates to an Early
Dynastic Period tomb in Saqqarah, Egypt circa 4900 BP (Helbaek cited in Solomon 1965). The
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granary weevil remains present in Saqqarah throughout the Early Dynastic Period and Old Kingdom
then becomes absent from archaeological contexts in the region for around 1000 years. Saqqarah is
believed to have been the necropolis for the ancient city of Memphis, which was constructed after the
unification of Upper and Lower Egypt. Traditionally, the unification is credited to Menes (Herodotus
HH II.XCIX 1791; Manetho 1940) as early as 5050 BP (Kitchen 1991; Savage 1998). Because of the
cultural link between the tombs at Saqqarah and Memphis and the alleged transportation of the cereals
to the tombs from the city as part of the funeral procession, the grain pests most likely infested the
cereal stores at the capital rather than having entered the context at the necropolis. The archaeological
disappearance of the granary weevil from Egypt also coincides with the relocation of the Egyptian
capital to Thebes at the start of the Middle Kingdom.
Sitophilus granarius may have entered the Egyptian contexts as a result of early trade with
Mesopotamia. Although the species has been recorded from Neolithic contexts in Israel (Kislev et al.
2004), the Sinai Peninsula would have presented a multitude of geographical obstacles (deserts, river
valleys, mountains, etc.) that would have inhibited, if not prevented, the granary weevil’s dispersal
into Egypt by the Bronze Age, without human assistance. Meanwhile, the recovery of Sumerian Late
Uruk and Jemdet Nasr Period (c. 5050- 4850 BP) pottery in Egyptian contexts (Roaf and Postgate
1991) supports the existence of a trade link between the regions. However, the resulting diffusion of
the granary weevil into Egypt only infers the transport of infested unprocessed grains from
Mesopotamia, not that the cereals were intended as a traded commodity. The grains could have
alternatively been used as a food source for the Mesopotamian merchants and/or their animals during
their journey to Egypt and have been deposited upon arrival in exchange for fresh cereals.
Archaeobotanical studies should be referred to in order to determine whether Mesopotamian cereals
were being imported into Egypt during the Early Dynastic Period.
The recovery of Stegobium paniceum, the biscuit beetle, from the Old Kingdom tomb of Queen
Ichetis at Saqqarah (Chaddick and Leek 1972) represents the earliest archaeological evidence of the
species. While the beetle is known to feed on a range of stored products (Buck 1958), it was found in
context with Sitophilus granarius suggesting a synanthropic association with cereals at this point. It is
unlikely that the biscuit beetle was endemic to Egypt as it has a preference for temperate environments
and a life cycle range similar to S. granarius [Figure 5.2]. If Stegobium paniceum was indigenous to
the Middle East, it could have been introduced into Egypt through a filter pathway across the Sinai or
anthropogenically via culture contact. Whereas the Sinai presented an obstacle for the unflighted
granary weevil, the biscuit beetle, capable of flight during its adult stage, may have been able to cross
the peninsula by dispersing between food sources (dried animal or vegetable material, Rees 2007). As
with Sitophilus granarius, Stegobium paniceum may have been introduced from the Middle East as a
result of trade and/or culture contact between the regions. Once introduced into Egypt, the biscuit
beetle maintains an archaeological presence through the New Kingdom Period. The species would
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have been better adapted for survival than Sitophilus granarius after the Old Kingdom Period society
collapsed because it is:
• a more generalist feeder, capable of surviving in the absence of cereals;
• a better disperser, permitting movement between settlements with or without human transportation;
and
• more tolerant of arid conditions (cf. Evans 1983; Rees 2007).
During the Middle Kingdom and New Kingdom Periods, the Egyptian contexts indicate the
introduction of insect species tolerant of warm temperate and tropical regions: Alphitobius diaperinus,
Palorus ratzeburgi, Rhyzopertha dominica, Tribolium castaneum, and Trogoderma granarium,.
These are species which have an optimal range for their cycle above 30 °C [Figure 5.2], and in the
case of T. granarium, require a minimum temperature of 24 °C for breeding (Hunter et al. 1973). The
continued presence of Stegobium paniceum, which places the maximum temperature around 34 °C, as
well as the lack of earlier fossil records suggests that new introductions are for the most part
heterochthonous and probably the result of the expanding Egyptian trade networks.
The particularly warm temperature requirements of Tribolium castaneum and Trogoderma
granarium pose an interesting problem by suggesting a more tropic region of origin, perhaps India,
despite earlier fossils of the species having not been found outside of Egypt. The granaries excavated
at Harappa and Mohendojaro (Wheeler 1966) would have been ideal habitats for the two species
(Buckland 1981), and Caspers (1972) provides an argument for coastal trade activity around the
Arabian Sea at the time. While step-wise trade could have occurred between Egypt and the Indus
Valley via the Fertile Crescent, the absence of the Tribolium castaneum and Trogoderma granarium in
the Middle Eastern sites would suggest a more direct form of trade between the two regions.
The life cycle requirements and optimal ranges (30-33 °C) of Alphitobius diaperinus and
Palorus ratzeburgi infer a warm temperate origin similar to modern-day Mediterranean Europe. The
species could have been introduced through culture contact with the Aegean kingdoms; however, the
species have not been recovered archaeoentomologically from contemporaneous sites in those regions.
Another option is the North African coast between Tunisia and Morocco. As archaeoentomological
investigations have not been conducted in the region, the Bronze Age insect fauna remains a mystery.
The lesser grain borer Rhyzopertha dominica is tolerant of both warm temperate and tropic
climates. Given the archaeoentomological evidence for potential Middle Kingdom connections with
India, it is possible that the species was introduced alongside the khapra beetle or Alfieri’s Tribolium
sp., if perceived as T. castaneum (cited in Andres 1931). However, the lesser grain borer does not
appear as thermophilic as either Trogoderma granarium or Tribolium castaneum, having an optimal
range around 33 °C inferring another origin. It is also probable that R. dominica was of African origin
from either the temperate North African coastline or tropical sub-Sahara. As stated above, there is not
a fossil record for the North African coastline; however, the lesser grain borer has been recorded in a
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Middle Bronze Age site in Spain (Alonso and Buxo 1993). If Marinval’s (1992) proposal for a North
African agricultural diffusion into Spain is considered, it could tentatively stand as evidence of R.
dominica’s presence in North Africa. The recovery of Cryptolestes turcicus (a species which has
optimal requirements similar to P. ratzeburgi and Alphitobius diaperinus) at Pharaonic Amarna
(Panagiotakopulu 2003) could reinforce the North African connection. The possibility of a subSaharan tropic origin for the species, and thus evidence of a trade connection for the Egyptians, is
largely speculative based upon the lesser grain borer’s life cycle requirements.
However, the
Pharaonic Amarnan presence of Tribolium confusum and Palorus subdepressus could also imply the
movement of cereals between sub-Saharan Africa and Egypt. Both species possess a life cycle similar
to Rhyzopertha dominica and are not as thermophilic as Trogoderma granarium and T. castaneum.
Moreover, P. subdepressus is widely distributed in the tropics in modern times (Rees 2007).
The archaeoentomological evidence supports the existence of a trade network between the
Egyptians and the Minoans. While the presence of koulouras suggests that the Minoans were capable
of storing large quantities of grain and may have been largely self-sufficient (Kieser 2005), the insect
evidence supports the importation of commodities from southern Europe (Greece), Anatolia, and
Egypt. Prior to the Late Minoan Period sites, Oryzaephilus sp. (likely O. surinamensis based on
context and species associations) had only been archaeologically visible from the Neolithic Mandalo
context in Macedonia; its recovery from Akrotiri (Panagiotakopulu and Buckland 1991) implies
maritime trade connection between the regions. Similiarly, the presence of S. granarius at Akrotiri
(Panagiotakopulu and Buckland 1991), Knossos (Jones 1984), and Kommos (Shaw and Shaw 1995)
suggests a link to the Near East, and the Rhyzopertha dominica, Stegobium paniceum, and Tribolium
confusum specimens imply a trade connection to Egypt.
Additionally, the introduction of O.
surinamensis (Alfieri 1931; Zacher 1934ab; Zacher 1937) to and the reappearance of Sitophilus
granarius (Panagiotakopulu 1999) in Egypt during this period, suggests that cereals were being
imported and exported from the Minoan settlements. This could mean that cereals were being used as
a form of currency in the Eastern Mediterranean into the Middle and Late Bronze Age.
Were the Aegean and Egyptian traders content with their Eastern Mediterranean connections,
or did they expand westwards in search of new cultures and commodities? The Late Bronze Age
records of the granary weevil from Italy and France cannot be viewed as confirmation of westward
exploration on the basis of biogeographical evidence alone. While Sitophilus granarius may have
been introduced to those regions through contact with the Eastern Mediterranean cultures during the
Bronze Age, sufficient archaeoentomological analysis has not been conducted on Neolithic sites to
discard the possibility of an earlier presence, particularly when the species was so well established in
Germany. However, the species has not been noted on Bronze Age sites in Germany, which suggests
that a Bronze Age introduction of the species from the north was unlikely. The Middle Bronze Age
evidence of Rhyzopertha dominica in Spain (Alonso and Buxo 1993) could stand as evidence of
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culture contact with the Eastern Mediterranean, but as stated above, it is likely to represent a
connection with the North African coast settlements.
The most conclusive archaeoentomological support for a westward movement during the
Bronze Age comes from the recovery of Stegobium paniceum in Britain (Osborne 1989; Robinson
1991), the earliest accounts of the species in the region. Despite numerous archaeoentomological
assessments having been undertaken in the United Kingdom, the biscuit beetle has only been noted on
two Bronze Age sites and in both cases as a single specimen, which suggests that the species was not
indigenous. The additional absence of species from Continental Europe during this era is interesting
as it seemingly denotes a direct connection with the Egyptian (the term used here is inclusive of the
North African settlements in Tunisia, Algeria, and Morocco) and Eastern Mediterranean kingdoms.
Due to the absence of other grain associated insects in the Bronze Age British contexts, Stegobium
paniceum was unlikely to have been introduced during the transportation of cereals, but anthropically
imported in association with another stored product.
The Bronze Age archaeoentomological accounts are indicative of the appearance and diffusion
of most of the major stored cereal pests of antiquity. Egypt, in particular, seemed to serve as a hub for
the movement of grains as made evident through appearance of a number of heterochthonous species
of varying temperature preferences. The influence of the Eastern Mediterranean trade networks
appears to have been far reaching with the insect evidence suggesting direct contact spanning from
India in the east to Britain in the west.
5.4.3 The Iron Age
The Pre-Roman Iron Age was a time of socio-economic turmoil. The Egyptian power that had
dominated the Bronze Age collapsed, conquered first by the Assyrians during the Third Intermediate,
and whose fate seemed to be largely dictated by whichever empire controlled the Near East—the
Assyrians, the Persians, the Macedonians, etc.
The Pre-Roman Iron Age also witnessed the
emergence of the Phoenicians as a Mediterranean maritime power.
Unfortunately, the
archaeoentomological record is meager for the period.
In the Near East, the granary weevil maintains a presence at Tel Arad (Hopf and Zachariae
1921) and Horbat Rosh Zayit (Kislev and Melamed 2000). Despite the absence of grain pest records
(due to the lack of archaeoentomological assessments in general) from the region during the Bronze
Age, the species most likely indicates a continuation of the Neolithic populations, and the Sitophilus
granarius specimens, noted in the Minoan and New Kingdom contexts, can be perceived as a
reflection of the species presence in the area between the Neolithic and the Iron Age. Additionally,
Alphitophagus bifasciatus and Tenebroides mauritanicus may also be representative of Neolithic
remnants as opposed to later introductions. All three species have an optimal temperature for the
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completion of their developmental cycle falling between 25 and 30 °C, possibly reflecting a similar
region of origin. While the cadelle and the granary weevil have been recorded together in Neolithic
contexts, A. bifasciatus has a very limited fossil record. However, the two-banded fungus beetle has
been recorded living in the wild in Israel (Chikatunov et al. 1997) suggesting an adaptation to an arid
climate. It is possible that the Near East hosted environments meeting the temperate preferences of
the cadelle and granary weevil as well as the arid conditions for A. bifasciatus [Figure 5.1]. Although
the Iron Age sites may have contained some indigenous pests, the presence of Oryzaephilus
surinamensis is indicative of an introduced species. As with the Bronze Age Egyptian records, the
Israeli specimen likely denotes an original introduction that dates to the Minoan Period connection to
Aegean and mainland Greece. Having adapted to a warm temperate climate, the species is tolerant of
warm temperatures [Figure 5.2] and would have flourished in an environment suited for S. granarius
and T. mauritanicus.
Similarly, the account of Sitophilus granarius in Okruglo, Croatia (Smith et al. 2006) does not
necessarily signify an Iron Age introduction. As with the Bronze Age record from Italy (Fasani 1976),
it could represent a population that was established during the Minoan Period or possibly from the
Neolithic, if the pottery impression from Servia is considered.
During this period, the earliest accounts of Trogoderma persicum and Sitophilus oryzae have
been documented by Chu and Wang (1975). Though it is worth noting their early presence outside of
the Mediterranean arena, neither species appears to have become widely dispersed until the Postmedieval period (e.g. Carrott et al. 1995b). However, a single record of S. oryzae has been reported
for the Roman era (Knörzer 1970) supporting the occurrence of exchange of commodities between the
Orient and the Mediterranean cultures.
The Iron Age stored product pest records from Spain and Britain are likely the result of culture
contact established by the Phoenicians.
In Spain, Perales (1984) records two new species, S.
granarius and Tribolium sp., in a context with R. dominica. The Phoenicians were a major seapower
from 3150 to 2750 BP, and its North African city-state Carthage flourished into the Roman Period
(Markoe 2000). Whereas R. dominica has been recovered from a Bronze Age context, the presence of
S. granarius and Tribolium sp. reflects a connection with the Eastern Mediterranean cities, possibly
Phoenicia. However, although Tribolium has not been recovered from Phoenicia, T. confusum and T.
castaneum could have been introduced there from Egypt during the Bronze Age. Alternatively, all
three species have been found in archaeological contexts from Bronze Age settlements in the Aegean
and in Egypt, which may have served as ports for their diffusion into Spain.
A Mediterranean maritime connection with the British Isles is also reflected by the insect
remains. As with the Bronze Age, Stegobium paniceum has been recovered an from Iron Age site in
England. While the biscuit beetle’s presence could be attributed to a residual population from the
Bronze Age, neither the earlier (Osborne 1989; Robinson 1991) nor the Iron Age (Chowne et al. 1986)
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records suggest the existence of the well-established populations evident in the later Roman period.
However, though unlikely, a derivative of the Bronze Age populations cannot be discarded solely on
the basis of a biogeographical survey and phylogenetic methods should be pursued (cf. King et al.
2009). As with the Bronze Age sites, the Iron Age Stegobium paniceum is unaccompanied by more
discernibly grain associated species implying that contact between the regions did not involve the
transport of large quantities of unprocessed cereals, which is supported by the Roman geographer
Strabo’s (SG) remarks on the tin trade between the Phoenicians and the inhabitants of the British Isles.
Leading up to the Late Iron Age, there is little archaeoentomological evidence regarding the
Roman Republic. The best evidence for the presence of grain-associated insects is indirectly derived
from Varro’s De Re Rustica, where it is possible to ascertain a few locations that had infestations of
curculio [see 5.3].
Although species level identification is problematic, Varro’s sites roughly
correspond with the archaeobiogeographical distribution of S. granarius from the earlier Iron Age
contexts.
5.4.4 The Roman Empire
The later Mediterranean Iron Age played host to the rise of the Imperium Romanum and the
furthest expansion of Roman control. It also affected a radical shift in the distribution of cereals
around the Mediterranean. During the Roman Republic, until the annexation of the North African
provinces, grains were produced on a small-scale by farmers, with their surplus transported to Rome
as taxation. After the annexation, the grain trade remained largely centralized, but immense quantities
of cereals were imported to Italian ports, such as Ostia, from Egypt and Africa for redistribution to
citizens; a socio-economic change which Yeo (1946) proposed would have severely undercut cereals
grown in Italy itself, resulting in a redirection of the local agricultural efforts towards other
commodities. The movement of cereals was also influenced by the Roman military. The Roman
Republic did not maintain a standing army, and therefore the legions were supplied only during
campaigns, and act, in the Middle and Late Republic, which involved the maintenance of lengthy
supply lines distributed from the centralized source with supplements in times of crisis directly from
allied regions (cf. Roth 1999). During the Imperium, Augustus restructured the Roman military to
create a widely dispersed standing army.
The garrisons were fed through local resources and
supplemented by supply lines. However, whereas during the Republic the provisions needed to be
transported first to a central location then redistributed, the Empire saw the movement of cereals to the
garrisons directly from the grain-producing provinces (Roth 1999). As a result of these changes, the
Roman Empire provided an outlet for the wide-spread dispersal of the grain-associated insect fauna.
In Central Europe, the first half of the first century AD was a period of conquest transitioning
into military occupation for the Roman legions (Bakels and Jacomet 2003). The archaeoentomological
remains provide evidence for the importation of cereals. The reappearance of Sitophilus granarius in
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the region, the first post-Neolithic record of the species, indicates that the imported grains were
unprocessed as the weevil does not typically infest milled materials. The presence of the granary
weevil in contexts with Oryzaephilus surinamensis (cf. Cymorek and Koch 1969; Koch 1970) is a
commonly seen association around the coastal Mediterranean region as early the Bronze Age (see
above), and Oryzaephilus sp. and S. granarius have established populations in France and Italy during
the first century AD, e.g. Ponel et al. 2000; Dal Monte 1956, respectively. However, the absence of
other grain fauna at the German and Netherland sites (specifically those species common to North
Africa and the Middle East during the Bronze and Iron Ages) infers an introduction of the pests, and
thus the grains, from southern Europe.
Moreover, as Oryzaephilus surinamensis has not been
recovered from Spain at this date, France and Italy seem promising hubs for the dispersal. Knörzer’s
(1970) record of S. oryzae is notable as the species is previously unknown from the Mediterranean
region and represents the earliest European record prior to the Post-Medieval Period. The presence of
the rice weevil, during the first century AD, signifies the importation of exotic (luxury) cereal
commodities to Neus-Novaesium, Germany that originated in the Far East (considering Chu and
Wang’s 1975 record from the Iron Age).
The Roman invasion and subsequent occupation of southern Britain in 43 AD are made evident
by the additional arrival of grain pests to the region. The Romano-British sites, 1 Poultry (Rowsome
2000) and 21 Saint Peters Street (King and Hall 2008; Appendix 2), provide evidence towards a
military introduction of the fauna as the sites are closely dated to the onset of the Roman occupation;
in particular, the 1 Poultry specimens, which pre-date the Boudican Revolt of 60 AD’s burning of the
twenty-year-old Roman commercial settlement of Londinium, present-day London (Rowsome 2000).
Mason (2003) estimates that 3,500 tonnes of grain would have been needed to supply the invading
troops during the first three months, not including the supplies needed to support the animals.
Although Britain had been exporting cereals to Roman provinces since Julius Caesar, the burden of
supporting the legionaries in addition to the local populations would have required imported
supplements until the garrisons were established. A single legion would have required over 2,800
tonnes of grain per year to support the troops (Mason 2003).
Based on the species present at these early sites, can the fauna be used to delineate the port of
origin for the military supply lines? Because of their proximity to Britain, the European hubs seem the
most likely suppliers of cereals during the invasion. The possibility of a European grain source is
supported by the recovery of Sitophilus granarius and Oryzaephilus surinamensis, which have a first
century AD, and earlier, presence in the grain-producing regions of Europe (modern-day Germany,
France, and Spain). However, Palorus ratzeburgi and Cryptolestes ferrugineus have not yet been
recovered from contemporaneous sites in those regions. If P. ratzeburgi and C. ferrugineus were
absent from Europe during this time, the two species would signify the existence of a longer supply
line. Moreover, as the species are secondary pests of stored products, they would not have been able
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to infest undamaged cereals at their source of origin, which would suggest the presence and likewise
simultaneous introduction of a primary pest such as S. granarius. As Palorus ratzeburgi’s only preRomano-British record comes from Egypt (and Oryzaephilus surinamensis and Sitophilus granarius
have also been recorded there in similarly dated contexts), cereals may have been shipped from the
Egyptian and/or North African markets, but as these accounts date to the Bronze Age, the argument is
tenuous.
The absence of a pre-Roman fossil record for Cryptolestes ferrugineus does not help clarify the
problem. While modern populations of C. ferrugineus are capable of breeding in a range of climates
(cf. Howe and Lefkovitch 1957), the species developmental optimal range for completion of its life
cycle is around 35 °C with a relative humidity of 90 % (Rees 2004).
Given its temperature
specifications, Cryptolestes ferrugineus likely entered the Roman Empire from two regions:
1.) the Mediterranean warm temperate zone, or
2.) a tropic zone, and its tolerance of warm temperatures and high humidity is similar in range to
Trogoderma granarium and Tribolium castaneum, both presumably from the Indian
subcontinent.
However, considering the extensive movement of cereals that occurred around the
Mediterranean and the exchange networks between the Mediterranean cultures, sub-Saharan Africa
and India that date back to the early Bronze Age, the lack of an earlier record for Cryptolestes
ferrugineus is suspicious if the species was endemic to those regions. Another possibility may be a
late introduction from the island of Trapobane (Sri Lanka). Pliny the Elder NH VI.XXII portrays the
first encounter between Rome and Trapobane in a meeting between the Emperor Claudius (41-54 AD)
and four ambassadors from the island, and discusses the size, fierceness, and ferocity of Sri Lankan
elephants in comparison to the Indian species as depicted in earlier accounts by Onesicratus, an
admiral of Alexander the Great. The establishment of a first century AD trade market with Sri Lanka
would have provided a pathway for the introduction of new stored product insects, such as
Cryptolestes ferrugineus.
Egyptian Red Sea trade, at the time, seemed concerned with the
establishment of elephant hunting stations (SG XVI.IV.VII, 1877; NH VI.XXXIV.CLXX-CLXXV,
1635). Hypothetically, if the Romans imported live Sri Lankan elephants into the Empire for
performances or warfare, the grain species may have been introduced into the Mediterranean region at
the Egyptian ports along the east coast of Africa, in association with the fodder needed to sustain the
elephants on the journey across the Indian Ocean and the Red Sea. The port of call in Egypt would
have limited and controlled C. ferrugineus’ initial infestation and dispersal. A major port, like
Alexandria, would have enabled the species to disperse into Italy and throughout the Empire from the
Italian distribution centers. However, a minor port along the East African coast would not have
necessarily affected a mass diffusion of the species. As the species does not appear to have become
widely established in Egypt by the second century (it is absent from Mons Claudianus,
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Panagiotakopulu and van der Veen 1997, though this could be attributed to unsuccessful competition
with its congener C. turcicus, see Grain Fauna above) or first century Herculaneum, Italy (Dal Monte
1956), the species may never have reached the Alexandrian or Italian ports in sufficient numbers.
Instead, it may have diffused across land via the trade caravans into the North African provinces and
then into Britain. A North African pathway would also explain the introduction of Palorus ratzeburgi
(another species absent from the Mons Claudianus site, Pangiotakopulu and van der Veen 1997).
While the Romano-British records of C. ferrugineus and P. ratzeburgi infer a non-European military
supply line, the presence of O. surinamensis and S. granarius allows for archaeoentomological
evidence of additional supply lines extending from European grain-producers. Moreover, it is possible
that multiple populations (denoting different ports of origin) of the saw-toothed grain beetle and the
granary weevil were introduced into Britain, but genetic analysis would need to be conducted (cf.
King et al. 2009).
As the Roman legions extended their conquest into northern England, the grain associated
insects were transported to the northern forts, e.g. Carlisle (Kenward et al. 2000; Kenward and Carrott
2006), Ribchester (Large et al. 1994; Buxton and Howard-Davis 2000), and York (Hall and Kenward
1976; Kenward and Williams 1979).
The diffusion of Sitophilus granarius, Oryzaephilus
surinamensis, Palorus ratzeburgi, and Cryptolestes ferrugineus following the military advancement
indicates that supply lines were maintained, to some extent, overland from southern Britain.
However, the introduction of Tenebrio obscurus, the dark mealworm, and Tenebroides
mauritanicus implies that supply lines were established and/or maintained through maritime sources as
well. Although its congener Tenebrio molitor, the yellow mealworm (a potential endemic to Britain
inferred from its early presence; cf. Howard et al. 1999; Smith and Howard 2004, and cold-temperate
adaptation [Figure 5.2]) has an early British presence, T. obscurus does not have a previous British
record. However, the contemporaneous presence of the species in France (Ponel et al. 2000) may
imply a supply line extending from France to the British frontier forts. The Roman introduction of T.
obscurus to England appears to have negatively impacted its congener T. molitor in Britain with the
invasive dark mealworm seemingly replacing the endemic species in the area by the end of the Roman
Period (Kenward in press). The ecological and thermal similarities between the two species suggest
that they may have experienced allopatric, or even peripatric, speciation as a result geographic
isolation. Furthermore, the two species are fairly morphologically distinct (see Mound 1989), which
implies a long period of separation, possibily in the range of millions of year (Kenward pers. com).
Both species seem adapted to the maritime temperate zone of Western Europe and occupy similar
niches, which would result in competition [Chapter 2]. As such, populations of the yellow mealworm
were probably initially limited to Britain and T. obscurus to the western coastal regions of Europe.
However, the dark mealworm, as an introduced species, would have had the advantage in Roman
Britain. The landscape alterations and changes in grain storage practices brought about by the Romans
145
would have initially, potentially, displaced the indigenous T. molitor from anthropic contexts and into
the natural environment, e.g. birds nests (Mound 1989). Moreover, if the introduction of T. obscurus
from France and Western Europe was in cereals, the dark mealworm would have been directly
imported into anthropic contexts and presenting as more archaeologically prevalent than its congener.
In areas with established populations of T. obscurus, the yellow mealworm would have difficulty
competing in order to establish itself in grain contexts. However, in the absence of, or the existence of
limited, populations of the dark mealworm, T. molitor may have been able to re-invade the anthropic
contexts, explaining its sporadic archaeological record.
Although Tenebroides mauritanicus could survive in temperate environs of France and may
have been imported to Britain alongside Tenebrio obscurus, the species lacks a Roman fossil record in
the region. Biogeographically, this suggests a supply line extending to the Eastern Mediterranean
where the cadelle was last archaeologically visible in the Iron Age contexts of Israel (Kislev and
Melamed 2000). A connection to the Eastern Mediterranean is also supported by the introduction of
Alphitobius diaperinus to Papcastle, Cumbria (Kenward and Allison 1995) as well as Tribolium
castaneum to second century York (Hall et al. 1980).
The importation of cereals appears to have continued throughout the Roman occupation of the
island. The third century AD account of Palorus subdepressus from Alcester, Warwickshire (Osborne
1971) implies a connection to the Egyptian provinces, as does the fourth century record of Tribolium
confusum and Cryptolestes turcicus at Towcester, Northamptonshire (Girling 1983).
However, during the second century AD, there is evidence to suggest that the Britain was
exporting cereals. Cryptolestes ferrugineus and Palorus ratzeburgi are present at the site of Amiens,
France in context with Sitophilus granarius and Oryzaephilus surinamensis (Yvinec 1997; Matterne et
al. 1998). C. ferrugineus and P. ratzeburgi archaeobiogeographically represent new species in the
region, and in addition to the presence of S. granarius and O. surinamensis may be indicative of an
exported Romano-British “fauna package”; a combination of ecologically interrelated stowaways that
flourished in Britain and accompanied the movement of cereals beyond its shores.
Moreover, Pals and Hakbijl’s (1992) assessment of the palaeoecological remains from a late
second century sunken ship in the Netherlands confirms that the grain fauna were transported with the
movement of cereals during the Roman Empire. While the authors suggest the loess region of
Belgium as the potential source for the grain (Pals and Hakbijl 1992, 298), the distinctive arthropod
fauna provides evidence towards another origin. While the presence of Cryptolestes ferrugineus,
Oryzaephilus surinamensis, Palorus ratzeburgi, and Sitophilus granarius hints at a Romano-British
origin, as mentioned above, the fauna had also been introduced to France by the second century.
However, the identification of Tenebrio molitor from the ship substantiates a British connection. The
Woerden specimen represents the earliest record of the yellow mealworm outside of Britain. Pals and
Hakbijl (1992) also report Mycetophagus quadriguttatus, a fungal feeder associated with foods. While
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not necessarily indicative of the presence of cereals, the beetle supports an argument for a British
origin. As with T. molitor, prior to this account, the species appears to have been to the limited to the
British Isles (e.g. Robinson 1979; Smith et al. 1999; Smith and Howard 2004). The recovery of
Alphitophagus bifasciatus is interesting as the species does not have an earlier connection to Britain.
However, the two-banded fungus beetle does not have an extensive fossil record and may have easily
been imported to England along with the other Romano-British Period introduced species, i.e. the
cadelle, the lesser mealworm beetle, or the rust-coloured flour beetle, with Eastern Mediterranean ties.
On the basis of biogeographical accounts, the fossil insect assemblage from Woerden appears strongly
indicative of a British origin for the cereals. Furthermore, the true benefit of the Woerden site and its
insect assemblage is that it demonstrates that grain-associated insects were transported long-distances
in cereals during the Roman Era.
5.5 Conclusion
Biogeography provides a tool enabling the examination of a range of human activities. By
investigating the archaeoentomological evidence for changes in the distribution of the grain species
over time, hypotheses may be formulated in regards to the past occurrence of human migration,
cultural contact, and trade.
While biogeography is capable of providing a time frame for the
introduction and diffusion of grain-associates, the approach suffers from limitations, and the resolution
of its results is ultimately dependant on the quality of the fossil record. When provided with a wellestablished fossil record, such as that in Britain, a biogeographical approach clearly provides evidence
for the introduction of cereal pests with the arrival of the Roman garrisons.
However, when
confronted with meagre evidence, similar to the present Neolithic record, the method can only offer
some insight into human activities and largely limits inferences to the realm of speculation.
The paucity of the archaeoentomological assessments outside of the United Kingdom, an issue
addressed by Buckland 1981, continues to pose problems for the completion of successful
biogeographical assessments, particularly in attempting to ascertain accurate shifts in species’
distributional ranges. For example, Sitophilus granarius has been recovered from Neolithic sites in
northwestern Europe. The next record of the species in the region comes from 8,000 years later during
the Roman occupation. Was the granary weevil truly absent from the northwest during the Bronze and
Iron Ages? Was the northwest a potential source for the diffusion of the species into Bronze and Iron
Age sites in southern Europe? When applied in isolation, biogeography is unable to address the
problem without a more refined fossil record, and it is only when material archaeological accounts are
consulted (the trade and cultural contact between Britain and northwestern Europe during the Bronze
and Iron Age, cf. Childe 1957, would have provide opportunity for the introduction any extant
147
populations of the granary weevil to Britain prior to the Roman Era) that the species’ absence from the
region between the Neolithic and Roman Periods can be inferred.
The major limitation of the biogeographical approach is that it is unable to discern multiple
introductions of a species as it does not distinguish between different populations of the same species.
During the early Roman conquest and occupation of Britain, the Roman garrisons appeared to have
been receiving cereal supplies from several provinces. As populations of Sitophilus granarius appear
around the Mediterranean, the species was likely transported to Britain from multiple regions on
several occasions. However, in the absence of other pests, a biogeographical investigation of S.
granarius would be blind to these introductions and, as such potential trade connections to regions
with early Roman settlements would be obscured. This is evidenced in areas like Spain (Moret and
Martin Cantarino 1996) and the Netherlands (Kuijper and Turner 1992) where S. granarius has been
the only grain pest recovered from sites dating to the early Roman period, which renders the species’
potential to delineate culture contact largely ineffective. However, the presence of other grain pest
species would provide additional clues towards discerning any potential trade connections, which
would enhance the efficacy of the biogeographical approach.
Regardless, the application of the biogeographical method towards grain-associated beetles has
an advantage over the strictly palaeoecological assessments [Chapter 4]. It moves beyond a site based
investigation of product-associations and autochthonous-allochthonous components in an effort to
holistically map species’ distributions in a spatial and temporal context.
This allows for the
consideration of a human-historic, rather than solely ecological, component in the investigations. The
limitations of the approach may be curbed by the incorporation of isotopic [Chapters 6 and 7] and/or
phylogenetic analyses [Chapter 8].
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Chapter 6
Stable Isotopes (δ2H, δ13C, and δ15N) from Beetles are Geographic Indicators of
the Origins of Cereals
149
6.1 Introduction
It has been demonstrated that the stable isotope composition of animal body tissues closely
corresponds to that of their diets (cf. Gannes et al. 1998), and in recent years, this concept has been
applied to the study of animal migrations (e.g. Hesslein et al. 1991; Best and Schell 1996; Hobson
1999; Hobson et al. 1999; Rubenstein and Hobson 2004). Stable isotopes of carbon and nitrogen have
been employed to delineate geographically distinct populations of animals on the premise that stable
isotope ratios in foodwebs differ regionally, providing naturally occurring signatures in organisms that
can be traced to origin (e.g. van der Merwe et al. 1990; Vogel et al. 1990; Alisauskas and Hobson
1993). Moreover, stable-hydrogen isotope measurements (δ2H) in several species of Neotropical
migratory songbirds (Hobson and Wassenaar 1997) and monarch butterflies Danaus plexippus
(Hobson et al. 1999) exhibited a geographic fingerprint permitting the deduction of the North
American origins of individuals of the respective species. These studies make use of the fact that
deuterium in rainfall is reflected in plants (Yapp and Epstein 1982), and subsequently, the stablehydrogen and carbon isotopic ratios are transferred through the foodweb by trophic-level consumers.
The present evaluation seeks to apply stable isotope methods towards discerning the origins of
modern cereals as made evident through chitin analysis of their associated beetle fauna.
In
archaeological investigations, grain beetles commonly serve as evidence for the presence of cereals at
sites, and in the absence of charred plant material, may stand as the sole indicators that cereals had
been present.
Although stable isotopes measurements of chitin have been used to address
palaeoecological issues (e.g. Miller et al. 1988; Motz 2000; Gröcke et al. 2006), the applications have
not been employed towards disentangling palaeoeconomic questions such as the origins of traded
perishable products in the past.
By determining the applicability of stable isotopes towards
unravelling the issue using modern materials, this study will hopefully serve as a platform to launch
future investigations involving insect fossils.
6.2 Materials and Methods
6.2.1 Laboratory Rearing Experiment
Adult populations of the granary weevil Sitophilus granarius, which were initially provided by
Central Science Laboratories in Yorkshire, UK, served as Generation 1 (G1) in the experiment. The
egg, larva, and pupa stages of each individual of Sitophilus granarius occur within a single cereal
kernel, and at 30 °C fully formed adults emerge 3-5 weeks after hatching and live 7 to 8 months
(Knopf 1980; Rees 2007). The adult weevils do not moult, and thus their chitin should provide a
material that is metabolically inert following the adults emergence from the grain. This is tested
during the experiment using weevils bred on British barley that were subsequently allowed to feed on
cereals from other geographic regions. Because the granary weevils mature in the interior of the
150
cereal kernels, the isotopic signature of their chitin should more closely correspond to the value of the
starchy layer (endosperm) of cereals than their exterior seed coat (brancoat).
This was briefly
examined during the present experiment.
The barley (Hordeum vulgare) and wheat (Triticum sp.) were locally-grown and purchased in
York, UK, and the oats (Avena sativa) were gathered from Macon, Mississippi.
These cereals
provided a control with a stable-isotope measurement from a known geographic region to which both
the δ2H of the growing season precipitation and the chitin isotopic values could be aligned.
Buckwheat (Fagopyrum esculentum), purchased in York but with an unknown geographic origin, were
used in blind tests to assess whether stable-hydrogen and carbon isotope measurements of beetle chitin
can be used to determine the geographic origins of the plants. Additionally, groups of granary weevils
were bred in tubes containing all four cereals to serve as a second blind test.
Groups of 15 weevils from G1 were placed in 50 x 25 mm tubes half-filled with individual
species of grains. A proportion of the remaining G1 weevils were frozen for isotopic analysis. The
tubes were held at 28 °C for 30 days to promote the oviposition of the granary weevils. Generation 2
adult weevils emerged in the tubes 30 days after the removal of G1. Fifty percent of the second
generation specimens (G2a) were immediately frozen for the isotope experiments, and the remainder
(G2b) were each transferred to separate tubes containing a second cereal type and kept at 28 °C for 30
days. G2b was then frozen for analysis.
6.2.2 Preparation of Chitin
The elytra were separated from the thorax and placed in 5 ml glass tubes. The samples were
rinsed in a solution of 2:1 dichloromethane: methanol in order to remove waxes. To eliminate
proteinaceous material, the elytra were then fully immersed in 2 ml 10 % NaOH, vortexed gently, and
incubated for 72 hours at 110 °C with gentle agitation (Tsao and Richards 1952; Miller et al. 1988).
The NaOH solution was formed using distilled water of known isotopic value. After incubation, the
resulting products were added to a spin column, spun through a filter at 10000 g for 1 minute, and the
collection tubes emptied. The products were re-suspended in the distilled water and spun at 6000 g for
1 minute. The collection tubes were emptied and the step repeated three times. The product was then
transferred to 2 ml Eppendorf Biopur tubes then dried overnight at 60 °C.
6.2.3 Removing Exchangeable Hydrogen
Because natural chitin contains some strongly absorbed water (Muzzarelli 1977) that can
isotopically exchange its hydrogen with ambient H2O, resulting in isotopic noise, Schimmelmann and
Miller (2002) investigated various strategies to eliminate or isotopically control the exchangeable
hydrogen from chitin.
The most economical approach discussed in the study involved the
equilibration of exchangeable hydrogen in chitin in H2O of known δ2H, succeeded by drying and high151
temperature pyrolitic liberation of H2 from the chitin in the presence of excess carbon (cf.
Schimmelmann 1991; Schimmelmann et al. 1993). A similar procedure was adopted in the present
investigation, but was modified to include the use of a standard (whale baleen) to further account for
the presence of exchangeable hydrogen in the chitin (see below). In an effort to limit exposure to
multiple sources of exchangeable hydrogen, the water (-47.43 ‰ Vienna Standard Mean Ocean Water
standard, VSMOW; Appendix 3F) employed for the NaOH solution and rinsing stages of the chitin
preparation was used for equilibration.
6.2.4 Stable Isotope Analysis
Hydrogen
The equilibrated chitin (1 mg) was weighed into silver capsules (5 x 8 mm). Additionally, the
starch-layers at the centres of the grain kernels were separated from the caryopses using a scalpel and
weighed into capsules. The filled capsules were left open for a period of 4 days to allow the
exchangeable hydrogen in the samples to fully equilibrate with the moisture in the laboratory air. The
stable-hydrogen isotope assays were performed at Iso-Analytical Limited Laboratories in Cheshire,
UK. The sample capsules were sealed just prior to analysis.
EA-IRMS (Elemental Analyser - Isotope Ratio Mass Spectrometry) was used for analysis with
the samples and references placed in silver capsules, sealed, and loaded into an auto-sampler. The
sealed capsules were then placed in a furnace at 1080 °C and thermally decomposed to H2 and CO.
Any trace water produced was removed by magnesium perchlorate, while, any trace CO2 was removed
using a Carbosorb™ trap. The H2 was resolved by a packed column gas chromatograph held at 45 °C.
The resultant chromatographic peak entered the ion source of the IRMS where it was ionised and
accelerated. Gas species of different mass were separated in a magnetic field then simultaneously
measured on a Faraday cup universal collector array. Masses 2 and 3 were monitored for H2.
The reference material used for δ2H analysis was IA-R002 (mineral oil, δ2HV-SMOW = -111.2
‰). IA-R002 has been calibrated against NBS-22 (mineral oil, δ2HV-SMOW = -118.5 ‰), an interlaboratory comparison standard distributed by the International Atomic Energy Agency [Appendix
3G].
Test samples of IA-R002 and IAEA-CH-7 (polyethylene foil, δ2HV-SMOW = +100.3 ‰) were
analysed along with the samples as quality control checks. Moreover, BWB-II (whale baleen) with a
known non-exchangeable δ2HV-SMOW value of -108 ± 4 ‰ was assayed within each batch of samples.
The capsules containing BWB II were treated identically to those containing samples during weighing
and equilibration with laboratory air. By using the measured δ2HV-SMOW value for BWB-II in each
batch, a simple correction for exchangeable hydrogen to the δ2HV-SMOW data was able to be applied.
The results for δ2H [Appendices 3D-3E] are expressed in per mil relative to VSMOW in Formula 6.1:
δ2HV-SMOW ‰ = [(RSample – RVSMOW) / (RVSMOW)] x 1000 ‰.
152
Replicates of samples and inter-comparison material yielded an external reproducibility of better than
± 1.90 ‰ for δ2H measurements.
Carbon and Nitrogen
EA-IRMS was likewise employed for stable-carbon analyses. The samples and reference
materials were weighed into tin capsules (0.5 mg), sealed, and then loaded into an automatic sampler
on a Europa Scientific Roboprep-CN sample preparation module. The materials were placed into a
furnace held at 1000 °C and combusted in the presence of oxygen. Moreover, the tin capsules flash
combusted, which raises the temperature around the sample to ~ 1700 °C. The combusted gases were
then rinsed in a helium stream over a combustion catalyst (Cr2O3), copper oxide wires (to oxidize
hydrocarbons), and silver wool to remove sulphur and halides. The resultant gases (N2, NOx, H2O, O2,
and CO2) were swept through a reduction stage of pure copper wires held at 600 °C, in order to
remove any oxygen and converted NOx species to N2. A magnesium perchlorate chemical trap was
utilized to remove water, and a Carbosorb trap removed CO2 during nitrogen-15 analysis. Nitrogen
and carbon dioxide were resolved with a packed column gas chromatograph held at an isothermal
temperature of 100 °C. The resultant chromatographic peak entered the ion source of the Europa
Scientific 20-20 IRMS, was ionised and accelerated. Gas species of different mass were separated in a
magnetic field then simultaneously measured on a Faraday cup universal collector array. For N2,
masses 28, 29, and 30 were monitored, and for CO2, masses 44, 45, and 46 were assessed.
The reference material [Appendix 3C] employed for δ13C and δ15N analysis was IA-R042
(powdered bovine liver, δ13CV-PDB = -21.60 ‰, δ15NAir = 7.65 ‰). IA-R042, a mixture of IA-R005
(beet sugar, δ13CV-PDB = -26.03 ‰) and IA-R045 (ammonium sulfate, δ15NAir = -4.71 ‰) and a
mixture of IA-R006 (cane sugar, δ13CV-PDB = -11.64 ‰) and IA-R046 (ammonium sulphate, δ15NAir =
22.04 ‰) were run as quality control check samples during the sample analysis. IA-R042 was
calibrated against and traceable to IAEA-CH-6 (sucrose, δ13CV-PDB = -10.43 ‰) and IAEA-N-1
(ammonium sulfate, δ15NAir = 0.40 ‰). IA-R005 and IA-R006 were calibrated against and traceable
to IAEA-CH-6. IA-R045 and IA-R046 were calibrated against and traceable to IAEA-N-1. IAEACH-6 and IAEA-N-1 are inter-laboratory comparison standards distributed by the International
Atomic Energy Agency (IAEA), Vienna. The results for δ13C were reported relative to the PDB
standard, and the δ15N was calibrated to atmospheric N2 [Appendices 3A, 3B, 3I]. Formula 6.2 shows
the expression of stable C and N isotope ratios as:
δ13C or δ15N = [(Rsample/Rstandard) - 1] X 1000,
where R is
13
C/12C or
15
N/14N for δ13C or δ15N , respectively.
Replicates of samples and inter-
comparison material yielded an external reproducibility of better than ±0.10 ‰ for δ13C and ±0.23 ‰
for δ15N measurements.
153
6.3 Results
Table 6.1 δ2H (‰) VSMOW of G2a weevils and associated cereals
δ2H (‰)
VSMOW
Chitin (N)
-90.81± 7 (6)
-114.61± 9 (6)
-101.39± 9 (4)
-84.7 (1)
Associated δ2H (‰)
Cereal
VSMOW
Starch
Barley
-96.21± 6
Buckwheat -127.69± 7
Oats
-103.16± 7
Wheat
-88.94± 3
Body-diet
Fractionation
+5.40
+13.08
+1.77
+4.24
δ2H (‰)
VSMOW
Seed coat
-59.60
-81.60
-51.00
-70.00
6.3.1 δ2H and δ13C Measurements from the Control Component
The data presented in Table 6.1 illustrate the intraspecific variation of δ2H exhibited by granary
weevils bred in the same host cereal as well as in host cereals grown in different geographic regions.
Within the test groups of the same host cereal, there was evidence of intraspecific variation
approximating ± 9‰ in the oats- and buckwheat-associated weevils. This exceeds the levels of ± 6‰
that have been recorded in previous control experiments (Miller 1984; Miller et al. 1988) and may
indicate the remaining presence of exchangeable hydrogen.
The elytra were strongly correlated with the isotopic signal of the endospermal starch layers of
their host plants [Figure 6.1; Figure 6.2]. The chitin was slightly enriched, averaging +6.1 ‰, in
deuterium compared to the starch, and the analyses indicated an approximate correlation of 0.90 for
the stable-carbon [Table 6.2] isotope measurements. The body-diet fractionation in the δ13C is similar
Table 6.2 δ13C (‰)V-PDB and δ15N (‰)Air of G2a Weevils and Associated Cereals
δ13C (‰)
Chitin (N)
-27.64± 0.45
(5)
-25.38± 0.90
(4)
-25.55± 0.98
(6)
-20.45± 0.36
(4)
Associated δ13C (‰)
Cereal
Starch
Barley
-28.26± 0.59
Body-diet
fractionation
+0.62
δ 15N (‰) δ15N(‰)
Starch
Chitin
1.50± 0.36 2.89±0.22
Buckwheat -24.71± 0.20
-0.67
0.46± 2.39 2.27±0.17
Oats
-26.31± 0.18
+0.76
2.08± 0.67 2.46±0.14
Wheat
-22.23± 0.40
+1.78
2.19± 0.61 3.23±0.22
to the range (± 0.5-1.1 ‰) reported in other investigations for trophic level variation (e.g. DeNiro and
Epstein 1978; Wada et al. 1987; Ostrom and Fry 1993; Michener and Schell 1994; Ostrom et al. 1997;
Bocherens and Drucker 2003). The enrichment of the deuterium in the chitin is indicative of a typical
trophic level increase (e.g. Birchall et al. 2005). However, this was not supported by the stablenitrogen [Table 6.2]. The chitin contained heavier nitrogen than the starch (with an average body-diet
fractionation of -1.16 ‰) and exhibited little correlation (R2= 0.42) with the isotopic nitrogen from
154
Figure 6.1 Correlation of δ2H in weevil chitin and starch of host cereal
0
-160
-140
-120
-100
-80
-60
-40
-20
-20
0
δ2Hstarch = 0.7616 δ2Hchitin - 18.666
-40
δ H Chitin
R2 = 0.9551
-60
2
-80
-100
-120
-140
2
barley
δ H Starch
buckwheat
-40.00
-30.00
-20.00
oats
-10.00
wheat
0.00
0.00
-5.00
13
13
δ Cstarch = 1.14 δ Cchitin + 4.07
-10.00
R = 0.90
-15.00
13
δ C Chitin
2
-20.00
-25.00
-30.00
13
δ C Starch
Figure 6.2 Relationship between the δ13C in Sitophilus granarius and host cereals
the starch [Figure 6.3]. This may imply that the chitin nitrogen was acquired from a source other than
the endospermal nitrogen. The absence of trophic level increase was also found elsewhere in the
laboratory samples of aphids bred on sorghum (body-diet difference of 0:0; Ostrom et al. 1997).
However, the nitrogen signal may reflect the use of dietary nitrogen in cuticular formation, with the
depleted δ15Nchitin being attributed to the process used during the removal of the wax layer. When the
analyses were replicated at the University of Bradford with the wax-layer preserved [Appendix 3E], a
trophic level change was made evident through the stable-nitrogen results. While sufficient material
was not available to re-test
15
N-wheat, the barley provided a δ15Nchitin+wax value of 6.64± 0.54, the
buckwheat indicated a δ15Nchitin+wax of 6.40± 0.18, and the oats expressed a δ15Nchitin+wax of 6.56± 0.06.
The deuterium from both the chitin and the starch were depleted in comparison to the seed coats,
155
exhibiting an average difference of -31.33 ‰ and -37.45 ‰, respectively. This result confirms the
granary weevil’s developmental association with the interior of the grains.
Figure 6.3 δ15N of chitin versus δ15N of starch
2.50
15
δ N Chitin
2.00
barley
1.50
buckwheat
1.00
oats
wheat
0.50
0.00
0.00
1.00
2.00
15
δ N Starch
3.00
4.00
15
15
δ Nstarch = 1.1918 δ NChitin - 1.6776
2
R = 0.422
Table 6.3 presents the results of the barley-bred (G2b) weevils that were transferred to
additional cereals in order to assess the metabolic stability of chitin in non-moulting beetles.
Table 6.3 δ2H (‰)VSMOW of G2b that
were bred in barley then transferred to a
second cereal
2
Sitophilus granarius specimens were allowed to mature in
barley before being transferred to containers with individual
δ H (‰)
VSMOW
Chitin (N)
-91.35± 5.35 (3)
Associated
Cereal
species of oats, buckwheat, and wheat. The G2b weevils
Barley Oats
beetles, -90.81± 7. In comparison to the G2abarley specimens,
-87.09± 3.65 (2)
Barley
Buckwheat
Barley Wheat
-91.30± 2.98 (3)
exhibited a D/H ratio within the range of the barley-bred G1
the G2boats presented a standard deviation of 0.38, the
G2bbuckwheat 2.63, and the G2bwheat 0.34. This suggests that
the chitinous hydrogen remains relatively inert in adult forms
of non-moulting beetles and reflects the hydrogen signal incorporated during development rather than
the dietary habits after emerging from the grains.
OIPC (The Online Isotopes in Precipitation Calculator) was consulted to predict the isotopic
value for the average meteoric precipitation of the host plant’s total growing season (δ2Hp) and the
predicted period of maximum tissue growth for the cereal kernels (δ2Hg) based upon geographic
perimeters (e.g. latitude; Bowen 2007) [Table 6.4]. In each case, both the δ2Hp and δ2Hg were
enriched in comparison to the δ2Hchitin, δ2Hstarch, or δ2Hseedcoat of materials from the corresponding sites
[Fig. 6.4].
Both δ2Hp and δ2Hseedcoat followed the geographic trend suggested by the Raleigh
distillation model with light hydrogen presenting as more abundant in higher latitudinal regions.
However, δ2Hchitin and δ2Hstarch were inversely proportionate to the predicted meteoric precipitation for
the host plant’s growing season, and showed the availability of relatively high levels of light hydrogen
at low latitudes. This may be a reflection of continentality as there is a notable difference in the
deuterium ratios of longitude 88.34˚ W (-100 ‰ δ2Hannualprecipitation) and 1.04˚ W (-56 ‰
δ2Hannualprecipitation) when tabulated for a constant latitude 53.57° N and altitude 11.9 m.
Moreover,
depleted deuterium values were reported by Hobson and colleagues (1999) for butterfly keratin
retrieved from Metairie, LA (-96 ‰ δ2H, latitude 29.58° N; longitude 90.09˚ W).
Between the intra-regional Yorkshire cereals, barley and wheat, there was isotopic discrepancy
in both the stable-carbon and hydrogen analyses. These may most likely be attributed to differences in
altitude where the plants were grown, relative humidity, and seasonality (winter wheat versus spring
barley). Previous studies have shown that
13
C (e.g. Körner et al. 1988; Körner et al. 1991) and 15N
(Körner and Diemer 1987; Körner et al. 1988) increase in plants in respect to altitude and relative
humidity. As the cereals were purchased commercially, the location in Yorkshire where the grains
were cultivated is unknown. However, the relative humidity can be calculated based on the local
Table 6.4 Precipitation values based on geographic perimeters
Location
(Associated
Cereal)
Yorkshire,
UK (Barley)
Macon, MS,
USA (Oats)
Yorkshire,
UK (Wheat)
Geographic data1
Growing
Season
Latitude
Longitude
53.57° N
1.04° W
Feb-Sept
33.06° N
88.34° W
Sept-April
53.57° N
1.04° W
Sept- July
Period
of Max
Tissue
Growth
for
Kernel
MayJuly
JanMarch
MarchMay
δ 2Hp2,3
δ2Hg2,4
-52.85
-55.33
-36.5
-43.33
-62.78
-64.66
1
Coordinates retrieved from Google Earth (Google Inc 2009)
Average δ2H values predicted through OIPC (Bowen 2007)
3 2
δ Hp = Average meteoric precipitation for total growing season of host plant
4 2
δ Hg = Average meteoric precipitation for period of max tissue growth for host plant kernel
2
growing season of the cereals. The winter wheat would have been subjected to an approximate RH of
80.6, and the spring barley an RH of 74.5, which may partially explain the enriched stable-carbon and
nitrogen values of the wheat. However, relative humidity is not the sole factor influencing the stablenitrogen and carbon. The Mississippian oats were cultivated during a period of relative humidity
approximating 88 but exhibit a lower isotopic composition than the wheat, which implies the effect of
additional factors such as altitude. Yorkshire displays a very complex series of altitudinal variations,
which would have had a significant impact on the isotopic composition of the cereals. For example,
the Vale of Pickering, for the most part, lies less than 30.5 m above sea-level whereas the North
Yorkshire Moors are almost entirely above 243.8 m, and in parts above 426.7 m, above sea level
(Darby and Maxwell 1978).
157
Figure 6.4 Relationship of average growing season precipitation (δ2Hp) with δ2Hseedcoat, δ2Hstarch, and δ2Hchitin
0
-80
-60
-40
-20
-20
0
-40
-60
barley
oats
wheat
Linear (Chitin)
Linear (Starch)
Linear (Seedcoat)
-80
-100
-120
2
δ Hp
Figure 6.5 Relationship of average precipitation for period of maximum tissue growth (δ2Hg) with δ2Hseedcoat,
δ2Hstarch, and δ2Hchitin
0
-80
-60
-40
-20
-20
0
-40
-60
-80
-100
barley
oats
wheat
Linear (Chitin)
Linear (Starch)
Linear (Seedcoat)
-120
2
δ Hg
The deuterium results indicated that the barley crops were raised at warmer temperatures than
the wheat. As the grains were cultivated within the same region, the inferred temperature difference
conveys seasonality. δ2H has been shown to vary up to 70 ‰ seasonally at a single site (White and
Gedzelman 1984). An increase in temperature affects the rate of evapo-transpiration that occurs
within the environment. In studies conducted on the affects of evapo-transpiration and temperature
upon the deuterium signal from leaves of high rooted plants (e.g. Roden and Ehleringer 1991), an
enrichment of the deuterium was observed as the light isotopes in water vapour escape the surface
more readily than the heavy isotopes (Craig and Gordon 1965). In the present experiment, this was
apparent to some degree in the seed coats, which would explain their enrichment over the starches.
However, the deuterium values in all the samples were more depleted than their respective predicted
meteoric precipitation. The presence of higher levels of light hydrogen may suggest that the seed
coats and interior starches were not influenced as strongly by the temperature and evapo-transpiration
factors that affected the rainwater deuterium.
Regardless, the enrichment of the seed coats in
158
comparison to the starches may be indicative of evapo-transpiratory impact. Furthermore, the δ2Hstarchseedcoat
fractionation may provide evidence of exposure levels to high temperatures, which result in an
increase in evapo-transpiratory processes. The fractionation increases with respect to temperature and
decreases with latitude, which is the inverse of the effect predicted by the Raleigh distillation model,
and similarly the stable-hydrogen composition of the endosperm reveals retention of light hydrogen
rather than the expected enrichment of deuterium, which is also reflected in the deuterium of the
chitin.
6.3.2 Mixed Cereal Blind Tests δ2H
Table 6.5 presents the δ2H results from the blind tests conducted on Sitophilus granarius bred
in mixed cereal samples. The isotope values mirror the range of measurements expressed by the four
component cereals when standard deviation was taken into consideration, and the results could be
roughly divided into four categories relative to the cereals [Table 6.6]. Because of the breadth of
standard deviation, there
Table 6.5 Blind Test: δ2H (‰) VSMOW of G2a
weevils bred in mixed cereals
Table 6.6 Categorisation of blind test results Based
on δ2HChitin range in control experiments
granarius
δ2H (‰)
VSMOW
Chitin
Barley
(-90.81± 7 δ2HChitin)
GK1
-88.71
Buckwheat
(-114.61± 9 δ2HChitin)
GK1; GK2;
GK4; GK5;
GK9
GK6; GK7;
GK8
GK2
-91.53
Oats
(-101.39± 9 δ2HChitin)
GK5; GK6;
GK7; GK8
Wheat
(-84.7 δ2HChitin)
GK3
G2a Sitophilus
GK3
-84.66
GK4
-89.55
GK5
-96.09
GK6
-108.54
GK7
-108.30
GK8
-101.41
GK9
-90.52
were overlaps in the ranges of the control values, particularly between buckwheat and oats, resulting
in the cross-categorising of specimens. During the laboratory-rearing stage, the weevils demonstrated
a preference towards the unprotected kernels of buckwheat and barley over the hull-concealed oat
159
grains, and as such, it is probable that the cross-listed specimens in the oats category should be
attributed to the other cereals. However, based solely on the δ2H values presented by the examination,
further distinction cannot be inferred.
6.3.3 Buckwheat Blind Tests
The isotopic measurements for the weevils from the buckwheat samples [Table 6.1; Table 6.2]
expressed a strong correlation between the chitin and the starch-layers of the grains. The stablecarbon and nitrogen suggest a geographic origin with a low relative humidity and low altitude. The
depleted deuterium composition, while typically indicative of high latitudinal regions, was similar in
range to the low latitude oats. Following the trend presented by the other cereals, a high light isotopichydrogen value in the buckwheat starch and chitin may reflect exposure to high temperatures and/or a
low latitude origin.
6.4 Discussion
A strong positive correlation was observed between the δ2H value of chitin and the δ2H value
of the starch of the host plant (R2= 0.96). This indicated that the grain weevil δ2H reflected the
deuterium signal of its diet prior to metamorphosing to its adult stage. A slight trophic-level increase
was noted in the positive body-diet fractionation ranging from +1.77 ‰ in oats to +13.08 ‰ in
buckwheat. The trophic level enrichment was also exhibited in the δ15N assays conducted at the
University of Bradford ranging from +3.75 ‰ in barley to +4.48 ‰ in oats. The amount of hydrogen
fractionation reported here is similar to other laboratory controlled experiments. Miller (1984) noted a
+10 ‰ enrichment in wheat flour-fed Tribolium molitor and Periplaneta americana fed on sucrosesupplemented Purina while Hobson et al.’s (1999) milkweed-bred Danaus plexippus only exhibited a
body-diet fractionation of + 1.5 ‰. In the present experiment, the body-diet fractionation for δ13C was
negligible. Additionally, the G2b trial demonstrated that the hydrogen composition in Sitophilus
granarius chitin remained metabolically inert following synthesis.
Both the carbon and non-
exchangeable hydrogen in chitin were primarily representative of dietary C-H. However, can they be
used as geographic indicators?
A myriad of investigations concerning plant stable-hydrogen have found a strong relationship
between δ2H signal of the source water and the δ2H of the plant (cf. White 1988; Zeigler 1988;
Flanagan and Ehleringer 1991; Hobson et al. 1999). Although the deuterium ratio of the source water
was unknown in the present analysis, the deuterium results recovered for the seed coats (R2= 0.9844)
and the starchy-layer (R2= 0.9928) when correlated against average growing season precipitation for
the period of maximum tissue growth (δ2Hg) [Figure 6.5] indicate a primarily rainwater-based water
source during the cultivation period. However, the starch displayed a non-unity slope of -0.6626
160
compared to the 0.8826 value of the seed coats for the regression with the water δ2H. These results
differ in comparison to the deuterium results for the seed coats (R2= 0.9625) and the starchy-layer
(R2= 0.9767) with a non-unity slope of 0.7034 and -0.5396 respectively, which were calculated for the
average meteoric precipitation for the total growing season (δ2Hp) [Figure 6.4]. The subtle
discrepancies between δ2Hp and δ2Hg in the seed coat and starchy-layer most likely reflect temporal
variations for the primary period of stable-hydrogen acquisition is the respective tissue layers. The
stronger correlation present in δ2Hg graph suggests the majority of deuterium in the cereal kernels was
assimilated during the period of maximum tissue growth rather than gradually over the entire growing
season of the plant.
In general, plant tissues have been found to be depleted in deuterium relative to source water
(Estep 1980; Hayes 2001), and the milkweed results presented in Hobson and associates’ (1999)
laboratory rearing experiment suggest that this discrepancy increases in respect to an increased
presence of heavy deuterium in the source water. Moreover, Flanagan et al. (1991) have shown a
difference between stem water isotopic composition and leaf water isotopic composition ranging from
-39 ‰ to -51 ‰ within plants from a single locale, and have proposed that the variation corresponded
to water-length pathway. Between the endospermal starch and the seed coat, the discrepancy ranged
from -18.94 ‰ in wheat to -46.09 ‰ in the barley from Yorkshire. Within the plant species, the
starch-seed coat deuterium fractionation theoretically reflects a difference in water usage. The seed
coats may have been supplied water from a source, which was on average, more enriched in deuterium
than the endosperm (perhaps leaf water that was enriched through transpiration; see White 1988,
Ziegler 1988, Flanagan and Ehleringer 1991).
While water-length and water-use pathways may have been a factor within the individual plant
species, the starch-seed coat variation, as mentioned above, also corresponded with differences in the
average temperature of the growing season between the wheat and barley. With a growing season of
September to July, winter wheat would have been exposed to an average temperature of 12.82 °C,
whereas spring barley (growing season March through September) would have been subjected to an
average temperature around 16.67 °C. This inference is further supported by the starch-seed coat
fractionation displayed by the oats of -52.16 during an average growing season temperature of 19.88
°C. Moreover, studies on bone collagen have suggested that relative humidity could account for
regional variation (Cormie et al. 1994), but as indicated by the depleted nitrogen and carbon in the
oats compared the wheat, the effects of relative humidity are not apparent through a 1:1 correlation.
Unfortunately, the full impact of physiological and environmental factors on plant deuterium still
requires further investigation.
Nevertheless, the strong linear relationship noted between meteoric precipitation for the period
of maximum tissue growth and both starch and seed coats was also present in chitin demonstrating that
161
the deuterium value of the water source controlled the isotopic composition of the cereals and the
grain pests [Figure 6.5]. A linear regression of the data yields the expression:
δ2Hchitin = -0.7871 δ2Hg – 135.15 [Formula 6.3]
with a correlation of 0.9934. When this linear regression was utilised to predict the isotopic signal for
the origin of the buckwheat samples, the average meteoric precipitation for the period of maxium
tissue growth was approximated to have a δ2Hg-buckwheat of -26.10 ‰.
If the average meteoric
precipitation for the total growing season is considered using Formula 6.4:
δ2Hchitin = -0.6363 δ2Hp – 124.57 [Formula 6.4],
the buckwheat seeds can be hypothesised to have originated from a region with an δ2Hp-buckwheat of 15.65 ‰.
According to the Raleigh distillation model, both predictions would back-trace the
buckwheat to a low-latitude region.
Figure 6.6 Relationship between δ2H and δ13C values of chitin from Sitophilus granarius
-30.00
-20.00
0
-200.00
-10.00
barley
buckwheat
oats
wheat
δ H Chitin
-40
-60
2
-80
-100
-120
-140
13
δ C Chitin
0
-200.00
0.50
1.00
1.50
2.00
barley
buckwheat
oats
-60
-80
2
δ H Chitin
-40
2.50
-100
wheat
-120
-140
15
δ N Chitin
Figure 6.7 Relationship between δ2H and δ15N values of chitin from Sitophilus granarius
However, examination of the results indicates that the deuterium of chitin becomes more
enriched in deuterium relative to meteoric water in higher latitudes compared to lower latitudes and in
colder months in respect to warmer months; a trend that is consistent with the deuterium of the starch
162
Figure 6.8 Relationship between δ13C and δ15N values of chitin
13
δ C Chitin
0.00
0.00
-5.00
0.50
1.00
1.50
2.00
2.50
barley
buckwheat
oats
wheat
-10.00
-15.00
-20.00
-25.00
-30.00
15
δ N Chitin
but in contradiction to the seed coat values. This implies that the chitin-water relationship may be
more complex than the straight line approximation indicates. The physiological and ecological factors
resulting in the internal discrepancies of the individual species of host plants also affect the δ2H of the
chitin. In order to effectively back-trace the geographic origins from chitin and account for multiple
influences, Rubenstein and Hobson (2004) and Bowen et al. (2005) have offered a non-linear
approach that has had success in tracing origins of wildlife. However, because of the parameters set
forth in this experiment, there is insufficient geographic and species data to successfully pursue a nonlinear interpolation approach.
Regardless, through comparison of the data collated from isotopic (δ13C, δ2H, and δ15N)
analyses of the chitin, it was possible to glean insight concerning the geographic details of the blindtested buckwheat’s region of origin. The similarities between the oats and the buckwheat apparent in
Figure 6.6 demonstrate that the buckwheat was more closely aligned with isotopic signal of the
Mississippian oats than either the Yorkshire barley or wheat. Its close orientation to the oats signal
(δ2H-δ13C) may be indicative of a low latitude origin. Furthermore, the δ13C- δ15N and δ2H - δ15N
[Figures 6.7 and 6.8] buckwheat values were severely depleted in relation to the other cereals, which
primarily reflects differences in altitude and relative humidity. The results predict a region of origin
for buckwheat of low altitude and high aridity.
Further information may be ascertained in Figure 6.9, which suggests a strong linear model
(R2= 0.9375) for predicting temperature (T) in relation to δ2Hchitin:
T= -0.2785 δ2Hchitin – 14.48 [Formula 6.5].
If other potentially influential factors (proximity to ocean, day-length, elevation, relative humidity,
etc.) are ignored, the regression approximation predicts an average growing season temperature of
163
17.43 °C for buckwheat based on an isotopic deuterium value of -114.61 ‰ for chitin. This places the
buckwheat in a region with much higher growing season temperatures than the other three cereals.
Figure 6.9 Relationship between average temperature (°C) for season of cereal tissue growth and δ2H values of
grain weevil chitin
Temperature °C
15
10
barley
oats
5
wheat
0
-105
-100
-95
-90
-85
-80
2
δ Hchitin
6.5 Summary
This experiment has shown that the stable hydrogen and carbon isotopic compositions in
granary weevils (Sitophilus granarius) are closely correlated with those of the endospermal starchlayer of their host cereals. While the isotopic signature of the host cereals is in turn controlled by the
local hydrology and climate, the isotopic composition of seed coat and starch within each host plant
indicates discrepancy in water-usage and water-pathways that deplete the deuterium of the source
water. Through the analysis of stable-carbon, hydrogen, and nitrogen from associated chitin, certain
details can be ascertained concerning the geographic origins of the buckwheat samples, suggesting a
region of low-latitude, low altitude, high aridity, and high temperature. The combined application of
multiple isotopes increases the resolution of isotopes as geographic indicators, and may prove
effective as a palaeoeconomic tool for discerning trade patterns.
164
Chapter 7
The Application of Isotopic Analyses towards Insect Remains: Modern and
Neolithic Case Studies
165
7.1 Introduction
Chapter 6 introduced the concept of insect fossils as palaeoeconomic indicators and, through
laboratory experiments, explored the isotopic relationship between geographically distinct cereals and
their respective granary pests. It was found that the stable-isotope signal of the dietary source is
locked into the chitin, presumably during chitin synthesis, which occurs when the exoskeleton is
formed during metamorphosis into the adult stage. In Chapter 7, carbon-13, deuterium, and nitrogen15 ratios are employed towards insect remains recovered from a modern reconstructed Anglo-Saxon
village and three German archaeological sites dated to the Linearbandkeramic Neolithic period. In the
present study, the stable-isotope signatures acquired from the chitin are assessed both independently
and collectively in order to attempt recognition of insect species which are associated with host
materials that are indigenous to the location of the site, foreign to the site but endemic to the nearby
hinterland regions, and foreign to the region.
7.2 Methods
The elytra were selected for isotopic analysis of the modern and archaeological insect
specimens. To isolate the chitin, the insect remains were rinsed in 2:1 dichloromethane: methanol
then immersed in 1 M NaOH for 24 hours at 110 °C. The stable-isotope analyses were conducted at
Iso-Analytical Limited Laboratories in Cheshire, UK using EA-IRMS.
All δ2H results are expressed in typical delta notation, in units per mil (‰), and normalised to
the VSMOW-VSLAP standard scale. The reference material used for hydrogen isotope analysis was
IA-R002 (mineral oil, δ2HV-SMOW = -111.2), which was calibrated against NBS-22 (mineral oil, δ2HVSMOW =
-118.5 ‰) distributed as an isotope reference standard by the IAEA. Samples of IAEA-CH-7
(polyethylene foil, δ2HV-SMOW = -100.3 ‰) were analysed along with the samples as quality control
checks [Appendix 4C]. In order to account for exchangeable hydrogen, whale baleen (BWII) was
analysed alongside the chitin. The exchangeable hydrogen was then accounted for using the Formula
7.1:
δ2Hchitin-corrected = measured δ2Hchitin (-108/BWBII measured value).
Carbon-13 results are calibrated to the Chicago Peedee Belemnite (PDB), and the Nitrogen-15
values were normalised to the standard atmospheric N2. The reference material employed for δ13C and
δ15N analysis was IA-R042 (powdered bovine liver, δ13CV-PDB = -21.60 ‰, δ15NAir = 7.65 ‰). IAR042, a mixture of IA-R005 (beet sugar, δ13CV-PDB = -26.03 ‰) and IA-R045 (ammonium sulfate,
δ15NAir = -4.71 ‰) and a mixture of IA-R006 (cane sugar, δ13CV-PDB = -11.64 ‰) and IA-R046
(ammonium sulfate, δ15NAir = 22.04 ‰) were run as quality controls during the analyses. The IAR042 was calibrated against and traceable to IAEA-CH-6 (sucrose, δ13CV-PDB = -10.43 ‰) and IAEAN-1 (ammonium sulfate, δ15NAir = 0.40 ‰). IA-R005 and IA-R006 were calibrated against and
166
traceable to IAEA-CH-6. IA-R045 and IA-R046 were calibrated against and traceable to IAEA-N-1
[Appendix 3C].
7.3 Case Study 1: West Stow, Sussex
7.3.1 Site Information
While notably an archaeological site of international significance West Stow presently hosts a
program for the experimental reconstruction of Anglo-Saxon buildings, initiated in 1973 (West 1985).
Six buildings have been constructed on the site using the traditional tools, materials, and techniques
thought to have been employed during the Anglo-Saxon period. In particular, West Stow tests West’s
re-interpretation of the Grubenhaus, or Sunken-Feature Building (SFB), featuring ground level
planked floors covering a ‘pit,’ which served as a storage area (West 1969, 1985; Tipper 2004). The
reconstruction is located at latitude 52.18° N; longitude 0.4° E, and is at an approximate altitude of 26
m above sea level.
7.3.2 Collection Methods
The beetles were collected from the reconstructed buildings using pitfall traps placed on 6th
June 2008 and recovered 3rd April 2009. The traps consisted of cat food tins, 70 mm in diameter, halffilled with sodium chloride solution with one drop of detergent, which served as a wetting agent so
that the insects were not trapped on the surface film. The traps were placed so that the neck of the tin
was level with the surrounding soil surface, and covered with a 15 cm square of expanded aluminium
mesh of approximately 1 cm aperture (cf. Kenward and Tipper 2008). Traps W7 and W19 were
placed under the floor at the base of the sunken feature of the Old House, a reconstruction of a twopost SFB (SFB 21) with a suspended floor above the sunken feature. The insect remains were
recorded using a low-power binocular microscope and the initial identifications were performed with
the assistance of H. Kenward.
7.3.3 The Fauna
Of the coleopteran fauna recovered from the pitfall traps, a limited range of species was
selected for isotopic analysis. The specimens were chosen based on:
• species biomass in pitfall traps,
• ecological associations,
• assumed trophic level, and
• prevalence in archaeoentomological contexts.
Anobium punctatum (Deg.), the furniture beetle or woodworm, is a serious pest of worked
wood, e.g. structural timbers and furniture (Palm 1959). The furniture beetle is primarily synanthropic
167
in Britain but has been recorded in the wild in dead trees (Duff 1993), particularly in association with
parts of the trees where branches and bark had been removed (Hickin 1968). Anobium punctatum is
commonly believed to infest only old dead wood—oak sapwood of at least 60 years and twenty years
for softwoods (Hakbijl 1989; Kenward and Hall 1995).
The carabid Carabus problematicus (Hbst.) is a predominantly predacious ground beetle,
which is commonly recovered in woodland, grassland, heathland, and moorland (Lindroth 1974;
Harde 1984; Duff 1993). Eyre (2000) includes members of Lumbricidae (earthworms), Enchytracidae
(e.g. potworms), Nematoda (roundworms), Diplopoda (millipedes), Acari (mites and ticks),
Collembola (springtails), Diptera (true flies), and Coleoptera (beetles) amongst C. problematicus’
prey. While primarily carnivorous, C. problematicus has been known to feed on fungi (Eyre 2000).
Catops nigricans (Spence) is a saprophagous species that benefits from dead and decaying
insects (Topp 1990). It has been recorded in wood pastures, woodland margins, mammal dens and
nests (Koch 1989a), as well as under leaf litter (Topp 1990; Duff 1993) and faggots (Donisthorpe
1939). Topp (1993) and Grist and Gurney (1997) have found that C. nigricans’ life cycle is seasonally
synchronized through photo-period, with it entering diapause during the summer, reproducing in the
autumn, and being most active during the winter.
Cryptophagus scutellatus (Newman) is the smallest European Cryptophagus species. The
beetle is believed to be mycetophagous, feeding on fungi, and has been noted in haystack and
vegetable refuse (Hinton 1945; Lindroth et al. 1973; Koch 1989b). It has also been recorded in
association with ants, Formica sp. (Palm 1959; Koch 1989b).
Ptinus fur (L.), the white-marked spider beetle, is a typical but non-obligate synanthrope. The
beetle is remarkably polyphagous and will infest a wide-range of animal and plant products (Zacher
1927; Dillon and Dillon 1972; Koch 1989a). The white-marked spider beetle is very common in
cereal stores (Mound 1989), and has also been recorded in birds’ nests, hives, and hay and straw waste
(Horion 1953).
The granary weevil, Sitophilus granarius (L.), is considered an obligate synanthrope. Unlike
the spider beetle, S. granarius is oligophagous, known to feed on a restricting range of food
substances, i.e. cereals. The granary weevil is discussed in more detail in Chapters 5 and 6. While
grains were not stored on site for consumption, Sitophilus granarius is assumed to have been
associated with the cereals present in the wheat thatch, indicating that the thatch material was stored
prior to its use.
7.3.4 Carbon Isotope Results
The application of
13
C ratios is a well-established technique in the study of past dietary
patterns and subsistence strategies (Katzenberg and Pfeiffer 2000). These isotopic signatures can vary
spatially based on differences in biogeochemical processes, which in turn are passed through the
168
foodweb and evidenced in the consumers (DeNiro and Epstein 1978). Table 7.1 presents the stablecarbon isotope ratios of chitin from the six species of Coleoptera from West Stow. The predicted
results range from -24.76 ‰ to -27.88 ‰.
The stable carbon isotopic fractionation provides information regarding the types of plants in
the foodweb, in terms of C3 or C4 plants. C3 are typically temperate plants, grasses, shrubs and trees,
Table 7.1 Carbon isotope ratios from West Stow
and include economically important crops such as
Species
δ 13C ‰ PDB
wheat, barley, and oats. C4 plants, such as
Sitophilus granarius
-25.16
Cryptophagus
-24.76
sorghum and millet, tend to be arid adapted plants
scutellatus
and grasses. C3 and C4 plants are isotopically
Ptinus fur
-27.29
Catops nigricans
-27.39
distinguishable through assess-ment of the related
Anobium punctatum
-27.88
stable carbon fractionation, which is associated
Carabus problematicus
-26.09
with metabolic pathway for carbon fixation during plant photosynthesis (Peterson and Fry 1987;
Tieszen and Boutton 1988). C3 plants tend to have stable carbon isotope ratios are -26 ‰, and C4
plants have more enriched δ13C values approximating -12 ‰ (Schoeninger 1995).
All of the West Stow specimens fell within the C3 category. The woodworm beetle exhibited
the most depleted stable-carbon isotopic values, and Cryptophagus scutellatus was the most enriched
in δ13C.
Ptinus fur and Catops nigricans produced δ13C values close to Anobium punctatum.
However, Sitophilus granarius and Carabus problematicus were slightly more enriched in δ13C than
the woodworm.
In terms of diet, the herbivorous and mycetophagous insects had stable-carbon isotope
signatures that were, in general, more enriched than the omnivorous and carnivorous species.
Anobium punctatum was an exception as the beetle had a δ13C approximating the carnivores and
omnivores. Table 7.2 shows the adjustment of stable-carbon isotopes to predict the isotopic signature
of the primary photosynthetic source. In order to calculate the approximate carbon signature for the
primary photosynthetic organism from the chitin, the herbivores were adjusted by one trophic level,
carnivores by two, and the omnivorous Ptinus fur was modified by one and a half trophic levels. The
following formulas were employed to calculate the isotopic signature for the primary photosynthetic
(∆13Ct):
Formula 7.2: ∆13Ct = δ13Ch – 0.62;
Formula 7.3: ∆13Ct = δ13Cc – (δ13Cc – δ13Cx) – 0.62;
Formula 7.4: ∆13Ct = δ13Co – [(δ13Co – δ13Cx + 0.62)/2] – 0.62,
where δ13Ch is the measured value for the chitin of the herbivorous beetles, δ13Cc is the determined
ratio for the chitin of the carnivorous beetles, δ13Cx is the approximated average of stable-carbon
isotope measurements in the assemblage, δ13Co is the observed ratio for omnivorous beetles, and 0.62
is the average fractionation tabulated from δ13Cbody-diet of grain-associated weevils in Chapter 6. The
169
∆13Ct for the carnivorous and omnivorous species is only an approximate value based on available
data; it must be noted that an unmeasured δ13C element inevitably comprises a proportion of their
actual diet. Thus the ∆13Ct formulated here simulates an artificial rather than a real trophic step
adjustment; however, it is used to clarify the results through limiting trophic level distortion.
Cryptophagus scutellatus proposes an interesting problem as a fungal-feeder. Fungi are
typically enriched in δ13C relative to their associated substrate, and the level of enrichment is
dependent upon the type of fungi.
Ectomycorrhizal species are known to be 1- 5 ‰ enriched
compared to their host foliage (Hobbie et al. 1999; Högberg et al. 1999; Kohzu et al. 1999); whereas
other species exhibit a 0- 2 ‰ enrichment of δ13C over associated carbohydrates (Gleixner et al. 1993;
Hobbie et al. 2003). Will et al. (1986) have recorded the enrichment as high as 7 ‰. Henn and
Chapela (2001) have found a fractionation effect that differentiates the ecological groups of
ectomycorrhizal and saprotrophic fungi. Ectomycorrhizal species exhibit a mean δ13C of -25.29 ‰,
and saprotrophic fungi have a mean of -22.14 ‰. Overlap exists between the ecological groups in the
-25.19 to -23.98 ‰ range. The stable- carbon isotopic value measured for the beetle Cryptophagus
scutellatus would have been affected by the isotopic signature of its diet. The δ13Cchitin falls in the
range of overlap for the two ecological groupings of fungi. As such, the dietary source cannot be
determined solely through the assessment of the carbon enrichment, and the stable-nitrogen signature
needs to be consulted.
If an average stable carbon isotope enrichment is taken (2.25 ‰), the
13
calculated ∆ Ct for Cryptophagus scutellatus becomes -27.63 ‰.
Table 7.2 Predicted carbon isotopic values of primary photosynthetics
Species
Trophic level
∆13Ct
Sitophilus granarius
Herbivore
-25.78
Cryptophagus scutellatus
Herbivore
Ptinus fur
Omnivore
-27.81
Catops nigricans
Carnivore
-27.08
Anobium punctatum
Herbivore
-28.5
Carabus problematicus
Carnivore
-27.08
1
1
-27.63
A fungal-feeding species
Through comparison of the ∆13Ct predictions, it was apparent that the West Stow site contained
materials from three different stable-carbon isotopic sources. The thatch-cereal material is represented
by the enriched signature from the Sitophilus granarius. The structural beams and worked wood are
likely signified by the depleted values from Anobium punctatum. The ∆13Ct from the remaining beetle
170
species may be an indication of isotopic values from plants in the local environment. Under that
assumption, the thatch would have originated in a geographic region with a higher altitude and the
structural material from a location with a slightly lower altitude than the West Stow site.
7.3.5 Nitrogen Isotope Results
Stable-nitrogen isotopes are often employed towards discerning trophic level (e.g. DeNiro and
Epstein 1981; Hobson 1990; Gu et al. 1996; O’Connell and Hedges 1999; Kelly 2000). However, as
mentioned in Chapter 6, nitrogen can be used as indicator of altitude and relative humidity. Table 7.3
shows the nitrogen-15 results for the West Stow Coleoptera. The δ15Nchitin values range from -1.25 ‰
to 21.10 ‰.
Table 7.3 Nitrogen isotope ratios from West Stow
Table 7.4 Predicted nitrogen isotope signatures
of the primary photosynthetics
15
Species
δ N ‰ Air
Species
Sitophilus granarius
Sitophilus granarius
1.02
4.02
Cryptophagus
scutellatus
Ptinus fur
21.10
Cryptophagus
scutellatus
Ptinus fur
18.10 ≥ 15Nt ≤ -2.9
2.36
6.86
Catops nigricans
Catops nigricans
Anobium punctatum
∆15Nt
3.91
-1.86
-2.09
Anobium punctatum
-4.86
Carabus problematicus
Carabus problematicus
-4.25
-1.25
While trophic level is evidenced by nitrogen-15 in beetle chitin, it is not as clearly defined as in
studies involving terrestrial vertebrates (e.g. Craig et al. 2009). The mycetophage Cryptophagus
scutellatus was the most enriched in δ15N, and the woodworm beetle Anobium punctatum exhibited the
most depleted nitrogen-15 values. The predatory ground beetle Carabus problematicus was also fairly
depleted in δ15N, but the other species fell in a range between 3.91 ‰ to 6.87 ‰. In general, the
carnivores showed lower nitrogen-15 signatures than the herbivores.
While shown to be slightly variable (~1- 6 ‰) in dietary experiments (Hilderbrand et al. 1996;
Hobson et al. 1996; Ambrose 2000; Bocherens and Drucker 2003), most models assume a 3 ‰ trophic
enrichment of
15
N. For simplicity, the 3 ‰ trophic enrichment was adopted here, and the modified
values to primary photosynthetic (∆15Nt) presented in Table 7.4. Additionally, the enriched isotopic
nitrogen-15 value for Cryptophagus scutellatus is evidence of a diet consisting of primarily
ectomycorrhizal fungus (cf. Henn and Chapela 2001). Unfortunately, the range of δ15N values in
171
ectomycorrhizal fungi is variable and the nitrogen signature could not be sufficiently adjusted beyond
15
Nchitin-diet fractionation. However, a tentative range is offered.
The ∆15Nt ranges from -4.86 ‰ to 2.36 ‰ (excluding C. scutellatus). The depleted values
evidenced by Carabus problematicus, Anobium punctatum, and Catops nigricans (cf. Hebert and
Wassannar 2001) may be indicative of primary photosynthetics originating in nitrogen poor soils. The
depleted ∆15Nt may indicate a separate geographic source from the more enriched nitrogen-15 value of
the cereals represented by Sitophilus granarius. Moreover, the enriched ratios evidenced by Catops
nigricans in relation to Carabus problematicus and Anobium punctatum may reflect seasonality. As
indicated by the Sitophilus granarius bred in winter wheat [see Chapter 6], the
15
N values are more
enriched in the colder season, and as Catops nigricans has been noted to reproduce in the autumn and
be most active during the winter, a comparatively enriched nitrogen-15 ratio is expected in respect to
the warmer season species. However, because of the number of geographic and environmental factors
influencing stable-nitrogen isotopic values, even at the primary photosynthetic level, insight into the
geographic origins of materials remains obscured when
15
N values are considered independently of
other isotopic assessments.
7.3.6 Hydrogen Isotope Results
Stable-hydrogen isotopic ratios are indicative of both trophic level (e.g. Birchall et al. 2005;
Reynard and Hedges 2007) and latitude (e.g. Meehan et al. 2004; Bowen et al. 2005).
Of the six coleopteran specimens from West
Table 7.5 Deuterium ratios from West Stow
Species
2
δ H‰
Stow, sufficient chitin material for deuterium
VSMOW
analysis was only available from four beetle
Sitophilus granarius
Cryptophagus scutellatus
species, and the results are presented in Table
-84.68
7.5. The predicted stable-carbon isotopic ratios
---
Ptinus fur
-97.58
ranged from -97.58 ‰ to -76.17 ‰. The annual
Catops nigricans
-84.69
meteoric precipitation (δ2Hp= -56 ‰) for West
Anobium punctatum
Carabus problematicus
Stow was calculated using OPIC (Bowen
---
2007).
-76.17
In vertebrates, Birchall et al. (2005) recorded an average stable-hydrogen isotope difference of
46 ‰ between carnivores and the combined herbivore and omnivore group. Similar findings were
reported by Reynard and Hedges (2007), who noted a 40 ‰ to 50 ‰ difference between the herbivore
and human groups. A 30 ‰ to 50 ‰ step was observed between the herbivore and the omnivore
groups, and from omnivores to humans, a 10 ‰ to 20 ‰ difference (Reynard and Hedges 2007). In
the West Stow specimens [Figure 7.1], the carnivore, Carabus problematicus, exhibited the most
172
Figure 7.1 Deuterium-chitin results versus the average deuterium ratio for annual precipitation at West Stow
0.00
-60
-50
-40
-30
-20
-10
0
-20.00
δ 2Hchitin ‰
-40.00
Herbivore
Omnivore
-60.00
Carnivores
-80.00
-100.00
-120.00
δ 2H ‰ of Average Annual Precipitation
enriched deuterium values. However, the herbivorous granary weevil predicted a more enriched
stable-hydrogen isotopic ratio than the omnivorous white-marked spider beetle.
Moreover, the
deuterium signature determined for Sitophilus granarius (-84.68 ‰) approximated the δ2Hchitin for
carnivorous Catops nigricans (-84.69 ‰), and only an 8.51 ‰ variation separated the granary weevil
from the ground beetle. As the polyphagous Ptinus fur expressed a trophic level omnivore-carnivore
step between 12.89 ‰ and 21.47 ‰ (similar in range to the omnivore-human step found by Reynard
and Hedges 2007) to Catops nigricans and Carabus problematicus, respectively, the deuterium value
for Sitophilus granarius does not follow the expected trend for a herbivore indigenous to the same
micro-population as the other beetles. The δ2H of an endemic herbivore, or herbivore bred on a
locally cultivated plant source, would be predicted to be at least a full trophic step below the
carnivores and a half step below the omnivores. Therefore, the granary weevil is likely representative
of an imported component to West Stow.
7.3.7 Comparison of Isotopic Assays
The application of isotopic analyses to the insect specimens recovered from West Stow
revealed the presence of autochthonous and allochthonous materials on the site.
In all three
assessments, the isotopic signatures procured from Sitophilus granarius did not follow the trend of the
other species in the assemblage and were generally more enriched than expected. Additionally, the
stable-carbon isotopic ratio from Anobium punctatum was slightly more depleted than the other
beetles; however, the nitrogen-15 values were similar to the predictions of Carabus problematicus and
Catops nigricans. In this section, the isotope results will be plotted together in an effort to glean
further information concerning the geographic origin of the material components represented by the
chitin [Figures 7.2- 7.5].
173
Figure 7.2 Carbon-13 and Nitrogen-15 plot of the insect remains from the Old House, West Stow. The herbivorous
(green) and mycetophagous (pink) species are indicated by squares. The omnivorous species is denoted by a yellow
triangle, and the carnivores are indicated by red diamonds
West Stow
13
δ Cchitin
-24.00
-5.00
-25.00
0.00
5.00
10.00
Sitophilus granarius
15.00
20.00
25.00
Herbivore
-26.00
Mycetophage
-27.00
Omnivore
Carnivore
-28.00
-29.00
15
δ Nchitin
The ground beetle Carabus problematicus is viewed here as representative of the local isotopic
signal at West Stow. This is based on the assumption that its diet consists of earthworms, slugs, etc.
(Eyre 2000) which are organisms that would have inhabited the local soil and vegetation. Similarly,
Catops nigricans, as a carrion-feeder, is assumed to be an indicator of the local signal. As carnivores,
the beetles are predicted to exhibit trophic level-enriched isotopic signatures relative to the local
herbivores and omnivores. The isotopic differences between the species likely reflect seasonality and
variation in diet.
Figure 7.3 Plot of ∆13Ct against ∆15Nt
-10.00
-5.00
-25.5
0.00
S. granarius 5.00
10.00
15.00
20.00
-26
C. problematicus
-27
C. nigricans
13
∆ Ct
-26.5
-27.5
P. fur
C. scutellatus
-28
A. punctatum-28.5
-29
15
∆ Nt
The isotopic results of Ptinus fur are enigmatic. The deuterium and carbon-13 ratios exhibited
by the species are indicative of the pattern established by the carnivores, but the nitrogen-15 signature
was comparatively enriched. As the deuterium result was a trophic step below the local carnivores,
the enriched stable-nitrogen does not support the adjustment expected for a trophic level change. As a
polyphagous beetle, the enrichment may be evidence of a diet supplemented by nitrogen-rich materials
such as fungi or moulds. However, as indicated by Cryptophagus scutellatus, a fungi-based diet
enriches the isotopic values of both the carbon and nitrogen.
174
Figure 7.4 Carbon-13 and deuterium plot of the insect remains from the Old House, West Stow. Sitophilus granarius
is represented by a green square. Ptinus fur is denoted by a yellow triangle, and Catops nigricans and Carabus
problematicus are indicated by diamonds, red and black respectively
West Stow
-28.00
-27.50
-27.00
-26.50
-26.00
0.00
-25.00
-24.50
-20.00
-25.50
δ Hchitin
-40.00
2
-60.00
-80.00
-100.00
Sitophilus granarius
Ptinus fur
Catops nigricans
Carabus problematicus
-120.00
13
δ Cchitin
Figure 7.5 Deuterium and Nitrogen-15 plot of the insect remains from the Old House, West Stow. Sitophilus
granarius is represented by a green square. Ptinus fur is denoted by a blue triangle, and Catops nigricans and
Carabus problematicus are indicated by diamonds, red and black respectively
West Stow
-5.00
-50.00
-1.00
-60.00
-3.00
1.00
3.00
5.00
7.00
9.00
δ Hchitin
Sitophilus granarius
-70.00
Ptinus fur
Catops nigricans
2
-80.00
Carabus problematicus
-90.00
-100.00
15
δ Nchitin
The data from Figure 7.2 suggests that the carbon and nitrogen isotopic ratios for Anobium
punctatum reflect the local signal because the woodworm appears a trophic step below the carnivorous
Catops nigricans.
However, the beetle presented a nitrogen-15 value similar to Carabus
problematicus and was more depleted in δ13C than the other species.
This suggests that the
woodworms’ associated dietary source (the structural timbers and floorboards in the houses) may not
have been indigenous to West Stow. When the isotopic ratio for the primary photosynthetic [Figure
7.3] is considered, the signature for Anobium punctatum is independent of the local feeders. This
suggests that some of the worked wood used in the reconstruction of the Old House was imported to
West Stow.
Like the single isotope comparisons, the multi-isotopic assessment for Sitophilus granarius
implies that the weevil fed on a diet, originating from a different geographic region than the other
species. S. granarius is a highly synanthropic species and has not been recorded in the wild in Britain.
Given the weevil’s strong association with stored cereals, the species’ isotopic signature is most likely
175
representative of the wheat cereals used in the roof thatch. The isotopic enrichment evidenced by the
granary weevil chitin is suggestive of a geographic region of higher altitude than West Stow.
7.3.8 Synopsis
For the West Stow site, the insect evidence permitted calculation of the local isotopic signal for
carbon, deuterium, and nitrogen. When trophic level effect was assumed, unusual patterns were noted
in Anobium punctatum, Cryptophagus scutellatus, and Sitophilus granarius. The isotopic enrichments
presented by C. scutellatus were attributed to its fungi-based diet. However, the variations present in
the woodworm and granary weevil are believed to signify the importation of materials to the site.
7.4 Case Study 2: Neolithic Germany
7.4.1 Site Information
The three archaeological sites date to the Neolithic Linearbandkeramic Period. Eythra is
located at latitude 51.13˚ N, longitude 12.18˚ E, and 67.9 m above sea level. The material selected for
isotopic analysis is associated with Well 2, c. 7269 BP. Insect remains were also retrieved from the
contemporaneous site of Plaußig, c. 7219 BP. Plaußig is located near Leipzig at latitude 51.23˚ N,
longitude 12.27˚ E, and altitude 115.5 m. The LBK site of Erkelenz-Kückhoven dates slightly later
than Eythra or Plaußig, circa 7040 BP. Erkelenz-Kückhoven is located near Colon at latitude 51.03˚
N, longitude 6.2˚ E, and 92 m above sea level.
7.4.2 Collection Methods
Environmental samples were selected from well contexts during the respective archaeological
excavations. The processing and the identification of the insect remains were conducted by E. Schmidt
at the Zoological Institute of the University of Freiburg using a binocular microscope and reference
collection. Official reports were generated for the sites: Erkelenz-Kückhoven (Schmidt 1998, 2010b);
Eythra (Schmidt 2005); and Plaußig (Schmidt 2010a).
7.4.3 The Fauna
From the identified faunal assemblages, Schmidt selected and donated beetle and fly
specimens for isotopic analysis.
The criteria established during selection of the West Stow
entomofauna were also used for the Neolithic Germany specimens.
The beetle Aphodius granarius (L.) is saprophagous, feeding on decaying matter, and
coprophilous, associated with excrement. The species appears to have a fairly polyphagous diet
(Landin 1961). Koch (1989b) has recorded the scarab in pastures, fields, and stables, especially in
176
rotting vegetation and herbivore dung. The larvae are believed to be predominantly saprophagous but
have been noted feeding on the roots of grasses (Hanski 1991).
The carabid Calathus fuscipes (Goeze) has been recorded in open habitats, particularly
cultivated fields and meadows (Lindroth 1974; Bengtson 1981; Luff 1988; Duff 1993), and under hay
and straw heaps (Koch 1989a). The species is purely carnivorous (Koch 1989a).
Carabus irregularis (F.) is a predatory ground beetle. Its head is disproportionate to the other
parts of its body and its mandibles are short and powerful (Casale et al. 1998). While little is known
about the prey of this species, the mandibles resemble those of Licinus specimens (species adapted to
opening shells), which suggests that C. irregularis may be a specialised snail hunter (Assmann et al.
2000).
Copris lunaris (L.) is a dung beetle that has been noted under herbivore excrement in
unploughed pastures with sandy soils (Jessop 1986; Koch 1989). The adults excavate a 10- 20 cm
tunnel under the dung leading to a brood chamber that usually contains 4 to 7 balls of dung holding
one egg each (Shirt 1987). The species is more commonly associated with cattle dung, but has been
found under sheep and horse manure (Koch 1989a).
The lesser stag beetle Dorcus parallelopipedus (L.) inhabits the rotten wood of broad-leaf
trees, especially the stumps (Donisthorpe 1939; Bullock 1993). The adults have been recorded under
the bark while the larvae develop within the trunk, branches, and roots (Palm 1959; Telnov n.d.).
Geotrupes vernalis (L.) is another beetle associated with dung. The species has been reported
in carnivore and herbivore excrement (Jessop 1986; Kuhne 1995) and has been recovered from bird
corpses (Jessop 1986). It burrows under the dung and lays one egg in each burrow (Jessop 1986).
Biström et al. (1991) have documented Hister (Atholus) corvinus (Germar) in dung and
decaying vegetation. It has also been noted in ant nests and the dens of foxes and sand martins
(Biström et al. 1991). The beetle is primarily predacious on maggots and other insects.
The dipteran Musca domestica (L.) is commonly known as the house fly. The larvae are often
present in nutrient-rich substrates such as faeces or decaying vegetation (Hewitt 1914; Amano 1985;
Hogsette 1996). Adults require a high protein diet in order to breed, and longevity is increased with
access to sugar (Lyske 1991, 1993).
Onthophagus ovatus (L.) (syn. O. joannae Goljan), like other scarabaeoid beetles, is primarily
coprophilous, and has been recorded beneath the dung of domesticated and wild animals (Koch 1989a;
Duff 1993), where it creates a pupal chamber in the earth (Whitehead 2006). The species may have a
rather polyphagous diet as Horion (1957) has described the species in carrion and rotting vegetation.
The chrysomelid Oreina caerulea (Olivier) is a leaf beetle associated with Cardueae
(Asteraceae) plant species (Pasteels et al. 1995). The beetle is purely phytophagous.
The Alfalfa snout beetle, Otiorhynchus ligustici (L.), is an occasional agricultural pest of
legume crops. The species is polyphagous but is often associated with the kidney vetch, Anthyllis
177
vulneraria (Shirt 1987; Bullock 1993; Morris 1997), and alfalfa, Medicago sativa (Vasilev 1914). The
larvae are root feeders (Shirt 1987).
Otiorhynchus raucus (F.) is a ground living weevil, which prefers chalky and sandy soils in
gardens and woodland (Hyman 1992; Morris 1997).
The commonly named root weevil is
phytophagous but feeds on a variety of plants. The larvae are root feeders while the adults frequent
the base of the plants and surrounding litter.
Pterostichus (Poecilus) cupreus (L.) is a thermophilic ground beetle commonly recorded on
arable land (Anderson et al. 2000). The species is predatory of cereal aphids and springtails (Kielty et
al. 1999; Mundy et al. 2000).
The pea leaf weevil Sitona lineatus (L.) is a capable of seriously damaging peas (especially
Pisum sativum), vetch (particularly Vicia faba), lentils, and fodder-beans (Jones and Jones 1974;
Bullock 1993; Morris 1997). Both the larvae and adults feed on the leaves and roots of leguminous
plants (Koch 1992). The weevil has been noted in gardens, meadows, and agricultural land (Morris
1997).
Sitophilus granarius (L.) and Tenebroides mauritanicus (L.) are synanthropic in cereals. As
detailed in Chapter 5, the cadelle is predacious (particularly on the larvae of Sitophilus and Stegobium;
Reitter 1911) but also attacks grains and cereal products (Koch 1989a). T. mauritanicus has been
recorded in the wild under tree bark of oak in Italy (Crowson 1958) and of beech and firs in Calabria
and Slavonia (Palm 1959).
7.4.4 Carbon Isotope Results
Table 7.6 presents the stable-carbon isotopic results for the three Neolithic archaeological sites. At
the site of Eythra, the δ13Cchitin values ranged from -24.62 ‰ to -18.28 ‰; the ground beetle
Pterostitus (Poecilus) cupreus gave the most depleted signature and the dung beetle Aphodius
granarius exhibited the most enriched stable-isotopic carbon. At Plaußig, the stable-carbon isotopic
values varied from -27.54 ‰ to -21.89 ‰. The root weevil Otiorhynchus raucus had the most
depleted results and Sitophilus granarius had the most enriched.
From the site of Erkelenz-
13
Kückhoven, δ Cchitin produced values extending from -29.38 ‰ to -23.30 ‰. The dung beetle Copris
lunaris had the most depleted isotopic value and Sitophilus granarius had the most enriched signature.
All of the isotopic predictions for the LBK sites were within the accepted range for C3 plants. Each of
the sites exhibited an isotopic fractionation of approximately 6 ‰. In order to clarify the results and
account for trophic level variability, Formulas 7.2- 7.4 were employed to calculate ∆13Ct [Table 7.7].
For Plaußig assessments, the isotopic ratio for Sitophilus granarius was excluded from the tabulation
of δ13Cx as the herbivore species was enriched compared to the rest of the assemblage which would
enrich the overall average. Additionally, because of the limited availability of materials, ∆13Ct could
not be formulated for the dung beetle from Eythra, and only a maximum value was predicted for the
178
Table 7.6 Predicted stable-carbon ratios from Neolithic Germany
Species
Calathus fuscipes
Onthophagus ovatus
Aphodius granarius
Hister (Atholus) corvinus
Otiorhynchus raucus
Sitophilus granarius
Aphodius granarius
Pterostichus (Poecilus)
cupreus
Sitophilus granarius
Musca domestica
Otiorhynchus ligustici
Geotrupes vernalis
Sitona lineatus
Copris lunaris
Dorcus parallelopipedus.
Oreina caerulea
Tenebroides mauritanicus
Sitophilus granarius
Carabus irregularius
δ13C ‰ PDB
Site
Plaußig
-23.39
Plaußig
-25.87
Plaußig
-24.74
Plaußig
-26.88
Plaußig
-27.54
Plaußig
-21.89
Eythra
-18.28
Eythra
-24.62
Eythra
-22.96
Erkelenz-Kückhoven
Erkelenz-Kückhoven
Erkelenz-Kückhoven
Erkelenz-Kückhoven
Erkelenz-Kückhoven
Erkelenz-Kückhoven
Erkelenz-Kückhoven
Erkelenz-Kückhoven
Erkelenz-Kückhoven
Erkelenz-Kückhoven
179
-25.23
-26.25
-28.20
-25.42
-29.38
-25.91
-29.37
-26.21
-23.30
-25.65
Table 7.7 Calculated carbon isotopic values of primary photosynthetics
Species
Calathus fuscipes
Onthophagus ovatus
Aphodius granarius
Hister (Atholus) corvinus
Otiorhynchus raucus
Sitophilus granarius
Aphodius granarius
Pterostichus (Poecilus)
cupreus
Sitophilus granarius
Musca domestica
Otiorhynchus ligustici
Geotrupes vernalis
Sitona lineatus
Copris lunaris
Dorcus parallelopipedus.
Oreina caerulea
Tenebroides mauritanicus
Sitophilus granarius
Carabus irregularius
1
∆13Ct
Trophic level
Carnivore
-28.16
Omnivore/ Herbivore1
-27.64/ -26.49
Omnivore/ Herbivore
1
-27.07/ -25.36
Carnivore
-28.16
Herbivore
-28.16
Herbivore
-22.51
Omnivore/ Herbivore1
Carnivore
-≤ -25.24
Herbivore
-23.58
Omnivore/ Herbivore1
-26.79/
-25.85
Herbivore
Omnivore/ Herbivore
1
-26.87
-28.28/ -28.82
1
-26.04
-28.87/ -30.00
Herbivore
Omnivore/ Herbivore
Herbivore
-26.53
-29.99
Herbivore
Carnivore/ Omnivore
-27.11/ -27.28
Herbivore
-23.92
-27.11
Carnivore
Associated with dung
ground beetle substituting δ13Cc for δ13Ch in Formula 7.2 as the carbon-13 signature of the Pterostichus
(Poecilus) cupreus was an unknown variable.
180
7.4.5 Nitrogen Isotope Results
Whereas it was possible to retrieve stable-carbon isotopic results from all the specimens,
nitrogen-15 assays could only be conducted on 15 of the 19 species, and Table 7.8 displays the results.
δ15Nchitin was only predicted for Sitophilus granarius from Eythra, and the granary weevil had an
stable-nitrogen isotopic signature of 6.89 ‰. From the Plaußig site, the four assessed species had a
range of 1.06 ‰ to 5.65 ‰. Onthophagus ovatus had the most depleted nitrogen-15 values and
Calathus fuscipes predicted the most enriched δ15Nchitin results. δ15Nchitin results were procured for nine
of the insect species from Erkelenz-Kückhoven, with a range of -6.02 ‰ to 10.10 ‰. The house fly
Musca domestica had the most depleted signature and the lesser stag beetle Dorcus parallelopipedus
had the most enriched values. In an effort to account for the trophic level variability within the
nitrogen isotopic results, ∆15Nt was predicted [Table 7.9].
Table 7.9 Tabulated nitrogen isotopic values of primary photosynthetics
Species
Calathus fuscipes
Onthophagus ovatus
Aphodius granarius
Sitophilus granarius
Sitophilus granarius
Musca domestica
Otiorhynchus ligustici
Geotrupes vernalis
Sitona lineatus
Copris lunaris
Dorcus parallelopipedus.
Oreina caerulea
Sitophilus granarius
Carabus irregularius
∆15Nt
Trophic Level
Carnivore
-0.15
Omnivore/ Herbivore
-3.44
Omnivore/ Herbivore
0.16
Herbivore
-0.27
3.89
Herbivore
Omnivore/ Herbivore1
-9.02
Herbivore
-5.16
Omnivore/ Herbivore
1
-8.99
Herbivore
-4.60
Omnivore/ Herbivore1
-2.22
Herbivore
7.10
Herbivore
0.09
Herbivore
-2.20
Carnivore
-7.66
181
7.4.6 Hydrogen Isotopic Results
Table 7.10 Deuterium ratios for Neolithic Germany
Species
δ 2H ‰ VSMOW
Site
Plaußig
Onthophagus ovatus
-119.98
Plaußig
Sitophilus granarius
-106.45
Eythra
Sitophilus granarius
-100.99
Erkelenz-Kückhoven
Musca domestica
-123.81
Erkelenz-Kückhoven
Otiorhynchus ligustici
-121.78
Erkelenz-Kückhoven
Geotrupes vernalis
-160.31
Erkelenz-Kückhoven
Copris lunaris
-130.46
Erkelenz-Kückhoven
Dorcus parallelopipedus.
-70.71
Erkelenz-Kückhoven
Carabus irregularius
-104.74
Sufficient archaeoentomological material to conduct stable-isotopic hydrogen assays was only
available from nine of the LBK specimens. Intra-site comparison was available to some extent;
however, because of the paucity of material, the values from three sites were consolidated to mimic a
bigger assemblage. The results were presented together in Table 7.10. For the three sites, the
Figure 7.6 Plot of δ2Hchitin against δ2Hp. Erkelenz-Kückhoven is represented by diamonds; Plaußig is indicated by
triangles; Eythra is denoted by a square. Herbivores are green, omnivore/ herbivore group is blue, and the
carnivore species is red.
-61.5
-61
-60.5
-60
-59.5
-59
-58.5
-58
-57.5
-57
0
-56.5
-20
-40
δ H chitin
-60
-80
2
-100
-120
-140
-160
-180
2
δ H Average Meteoric Precipitation
deuterium values varied over a range of -160.31 ‰ to -70.71 ‰. Geotrupes vernalis recovered from
Erkelenz-Kückhoven predicted the most depleted δ2Hchitin whereas the Dorcus parallelopipedus from
Erkelenz-Kückhoven exhibited the most enriched stable-hydrogen isotopic signature. The modern
182
annual meteoric precipitation for each of the three sites was calculated using OPIC (Bowen 2007):
δ2Hp- Eythra= -60 ‰, δ2Hp- Plaußig = -61 ‰, and δ2Hp- Erkelenz-Kückhoven = -57 ‰ [Figure 7.6].
7.4.7 Comparison of Isotopic Analyses and Discussion
The isotopic values were variable within and between the sites [Figures 7.7- 7.10]. The
herbivorous species, in particular, evidenced a large range of variation, which suggests an
allochthonous element for their associated plant species. This was especially apparent in δ13C, δ2H,
and δ15N results from Erkelenz-Kückhoven.
Figure 7.7 Carbon-13 and Nitrogen-15 plot of the insect remains from LBK Germany
Neolithic Germany
-5.00
δ13C
-10.00
0.00
0.00
-10.00
5.00
10.00
15.00
PlauBig
Erkelenz-Kückhoven
Eythra
-20.00
-30.00
-40.00
15
δ N
The dung fauna
The dung-associated species recovered from the Linearbandkeramic sites most likely reflect
the diet of the inhabitants, human and animal, living at the sites. The species, including the house fly,
lay their eggs directly in the manure or in dung balls manufactured from the excrement. When the
larvae emerge, they subsist on the nutrients and proteins available in the manure, which in turn are
used to help form the chitin after the species pupate.
Figure 7.8 Plot of ∆13Ct against ∆15Nt
-10.00
-8.00
-6.00
-4.00
-14
0.00
-16
-2.00
2.00
4.00
6.00
8.00
-18
∆13Ct
-20
Plaußig
Eythra
-22
-24
Erkelenz-Kückhoven
-26
-28
-30
-32
15
∆ Nt
183
At Eythra, Aphodius granarius exhibited an enriched δ13C value compared to the other species
from the assemblage that were analysed. The -18.28 ‰ ratio approaches the range expected for C4
plants (Schoeninger 1995). However, because of the period and location, the isotopic signature is
probably connected to an omnivorous animal or humans as the carbon-13 value from A. granarius is
4.68 ‰ more enriched than the signature from the granary weevil, approximating a trophic level step.
An herbivorous vertebrate such as a cow or a sheep would be expected to give an isotopic ratio similar
to Sitophilus granarius, if bred on a diet consisting entirely of cereals. An herbivore bred on a mixed
diet of cereals and local grasses would have an isotopic signature that is a blend of the two dietary
values. For example, if the ∆13Ct from the Pterostichus (Poecilus) cupreus is a reflection of the
carbon-13 ratio from the primary photosynthetic in the local environment (≤ -25.24), the herbivore
would be expected to predict an isotopic value in the range of -22.96 ‰ ≥ herbivore ≤ -25.24 ‰.
Alternatively, if the Aphodius granarius is considered to reflect the isotopic ratio transferred through
the dung of an indigenous herbivore, the local carbon-13 value would approximate -18.28 ‰, and the
granary weevil and ground beetle would reflect allochthonous materials.
Unfortunately, more
specimens were not available for analysis, and an association with omnivorous or higher trophic level
species is inferred for the dung beetle.
Figure 7.9 Plot of Deuterium and Nitrogen-15 from Neolithic Germany
Neolithic Germany
-8.00
-6.00
-4.00
0.00
-2.00 -20.000.00
2.00
4.00
6.00
8.00
10.00
12.00
-40.00
δ2 H
-60.00
PlauBig
Eythra
-80.00
-100.00
Erkelenz-Kückhoven
-120.00
-140.00
-160.00
-180.00
δ15N
The dung beetles from Plaußig may reflect an herbivore which had a diet consisting of the
local flora and cereals. The δ13C ratios for Onthophagus ovatus and Aphodius granarius fall close to
the carbon-13 signature for the root weevil Otiorhynchus raucus but below the δ13C ratio for
Sitophilus granarius. However, both the δ13C and the ∆13Ct values for the granary weevil fall outside
the average predicted isotopic range of the rest of the assemblage (∆13Ct-average = -27.76) suggesting
that the species may in itself be associated with allochthonous plants; whereas the ∆13Ct of the dung
beetles approximates the average for the assemblage. The addition of the deuterium and nitrogen-15
isotopes did not help to further elucidate the dung beetle’s association due the availability of relatively
few samples. The 1.13 ‰ difference in δ13C ratio between Onthophagus ovatus and Aphodius
184
granarius presents as a 3.5 ‰ variation in δ15N values. The variability evidenced by the isotopic
ratios of the dung beetles could signify differences in altitude of the associated herbivores’ grazing
ground or seasonality in the beetle’s chitin formation.
Copris lunaris, Geotrupes vernalis, and Musca domestica represent the dung fauna from
Erkelenz-Kückhoven.
The isotopic ratios were highly variable.
G. vernalis and M. domestica
exhibited depleted nitrogen-15 ratios while C. lunaris had a relatively high value for the assemblage.
Geotrupes vernalis was approximately 30 ‰ more depleted in deuterium than Copris lunaris and 37
‰ more depleted than Musca domestica. Copris lunaris exhibited the most depleted carbon-13
signature with Geotrupes vernalis approximately 1 ‰ more enriched and Musca domestica about 4‰.
Figure 7.10 Plot of Deuterium and Carbon-13 from the German palaeoentomological specimens
Neolithic Germany
-30.00
-25.00
-20.00
-15.00
δ2 H
-35.00
-10.00
0.00
-5.00
0.00
-50.00
-100.00
-150.00
PlauBig
Eythra
Erkelenz-Kückhoven
-200.00
13
δ C
The discrepancy among the dung fauna can probably be attributed to differences in grazing areas and
seasonality. The trophic level of the manure source may have been a factor, but the isotopic assays
gave conflicting results. For example, the δ15N ratio of the house fly was more nitrogen poor
(implying a lower trophic level position) than expected relative to its deuterium signature. Despite the
interspecific variation, the three dung-associates appear to be representative of herbivorous dung
sources. The deuterium and carbon-13 values of the house fly are similar to those predicted for the
Alfalfa snout beetle Otiorhynchus ligustici [Figure 7.10] and the carbon-13 and nitrogen-15 values of
Copris lunaris are grouped close to the chrysomelid [Figure 7.7]. The stable-carbon isotope value for
Geotrupes vernalis is also similar to Oreina caerulea but the dung beetle’s nitrogen-15 prediction is
more depleted.
The depleted stable-carbon and deuterium values (relative to Musca domestica,
Carabus irregularis, Sitona lineatus, and Otiorhynchus ligustici) of Copris lunaris and Geotrupes
vernalis suggest the exploitation of a lower altitude areas for the grazing of herbivores.
While previous studies have shown that manure increases the
15
N content of soils and plants
(e.g. Choi et al. 2003; Bogaard et al. 2007), the chitin of the dung fauna did not appear to exhibit any
nitrogen-15 enrichment. In comparison to the other species in the assemblages, the dung-associates, in
general, were more nitrogen deprived. This suggests that the majority of the dung associated insects
were not feeding directly on the nitrogen-enriched aspects of the dung. However, Copris lunaris may
185
be an exception as it was slightly enriched relative to the majority of the invertebrate assemblage from
Erkelenz-Kückhoven.
The plant-associated insect species
The invertebrate remains from the Neolithic German sites may give evidence concerning the
exploitation of natural resources, agricultural practices, and possibly trade. As discussed in Chapter 4,
insects are very important indicators of plant resources (e.g. wood, hay, cereals, dyeplants, moss, turf
and brushwood).
While the traditional archaeoentomological evaluations permit inferences
concerning the probable origins of the plant resources, the application of isotopic analyses may be
utilised to substantiate the otherwise largely hypothetical palaeoecological suggestions.
Only a limited range of plant-associated species were available for isotopic analyses from the
Early Linearbandkeramic sites of Eythra and Plaußig. The granary weevil represents the Eythra
assemblage and Sitophilus granarius and Otiorhynchus raucus were selected for isotopic evaluation
from Plaußig. At Eythra, the granary weevil was more enriched in carbon-13 than the predatory
Pterostichus (Poecilus) cupreus but more depleted than the dung beetle.
Based on the assumption
that δ13C ratios are more enriched in carnivores than herbivores within the same foodweb, P. cupreus
was not feeding on S. granarius, and the two species likely reflect the isotopic values of separate
areas. The granary weevil would likely represent cereal cultivated at a higher altitude or lower latitude
than occupied by the ground beetle’s prey.
While both Eythra and Plaußig were discovered in the Leipzig region of modern Germany, the
granary weevil specimens from the two sites predicted different isotopic values. The Sitophilus
granarius recovered from Eythra had more enriched stable-nitrogen and deuterium isotopic ratios but
more depleted stable-carbon values than the Plaußig specimen. As Plaußig is located at a higher
altitude than Eythra but approximately the same latitude, the isotopic signatures of the local flora and
fauna should be more enriched in the Plaußig specimens. The intraspecific variation may reflect
differences in seasonality, nitrogen-content in the soil, or cereal species.
When the root weevil Otiorhynchus raucus is considered, the stable-carbon isotope values for
the plant-associated fauna at Plaußig vary by approximately 6 ‰. As the stable-carbon isotopic value
for the root weevil approximates the average ∆13Ct of the rest of the assemblage, the species is most
likely autochthonous, which would imply that the δ13C ratio for the local environment around Plaußig
is probably close to -27 ‰. Based on difference in elevation, the local environment signature for
Eythra would be ≤ -27 ‰, which supports the prediction of ≤ -25.24 ‰ tabulated from Pterostichus
(Poecilus) cupreus. This would suggest that the granary weevil specimens from Eythra and Plaußig
reflect allochthonous cereals. While the enriched isotopic values of the granary weevils may signify
the importation of cereals to the sites from slightly lower latitude regions, the enrichment most likely
supports a local cultivation of the crops in the higher altitude hinterlands (see Körner et al. 1991).
186
Both sites are at a lower elevation than their surrounding hinterlands. In comparison to Eythra and
Plaußig, more plant-associated invertebrates were available for analysis from the later
Linearbandkeramic site of Erkelenz-Kückhoven: Dorcus parallelopipedus, Oriena caerulea,
Otiorhynchus ligustici, Sitona lineatus, Sitophilus granarius, and Tenebroides mauritanicus.
As mentioned above, the recorded isotopic values for Oriena caerulea, Otiorhynchus ligustici,
and Sitona lineatus, though variable amongst themselves, are similar to the dung fauna. The species
may have been transported to the site as the Neolithic people exploited their local and hinterland plant
resources. O. ligustici and S. lineatus, in particular, are associated today with edible plant species.
Both the pea leaf weevil and the Alfalfa snout beetle exhibited similar isotopic predictions to the
ground beetle Carabus irregularis. It is likely that the beetles represent the isotopic signature for the
local vegetation. Given the isotopic correlation with Musca domestica, the house fly was probably
breeding in dung and waste materials on the site.
The depleted stable-carbon ratio of Oriena caerulea may reflect the use of a lower altitude
region compared the pea leaf weevil and the Alfalfa snout beetle. As the hinterland decreases in
elevation to the east of Erkelenz-Kückhoven, this may be a likely origin for the species. The nitrogenenrichment evidenced by O. caerulea in addition to its stable-carbon ratio similarity to Copris lunaris
and Geotrupes vernalis may imply that the chrysomelid originated from the region set aside for the
grazing of the domesticated vertebrate herbivores. As Choi et al. (2003) and Bogaard et al. (2007)
have shown, manure causes nitrogen enrichment in plants. While probably not intentionally fertilized
by the Neolithic people, the Cardueae (Asteraceae) plants would have indirectly benefited from the
presence of herbivore dung. The enriched nitrogen values would have then been transferred to the
phytophagous Oriena caerulea.
The isotopic values predicted for Dorcus parallelopipedus are interesting. The lesser stag
beetle exhibited a stable-carbon isotopic ratio similar to the perceived autochthonous insect species;
however, had much more enriched δ2H and δ15N values. The deuterium signature of the stag beetle is
approximately 51 ‰ more enriched relative to Otiorhynchus ligustici and the nitrogen-15 fractionation
around 12 ‰. As both beetles are herbivorous, the discrepancy suggests the introduction of D.
parallelopipedus to Erkelenz-Kückhoven. The lesser stag beetle is often associated with rotten or old
wood, and in a village, its habitat would most likely be mimicked by the presence of wood piles
intended for firewood. While Dorcus parallelopipedus may have been transported to the site along
with the wood, the isotopic discrepancy presented by its chitin suggests that the wood would have had
to have been transported a fair distance from a warmer region. As the beetle is flighted, it is more
likely that the species originated in a warmer environment and arrived at the site through natural
means rather than anthropic.
Both Sitophilus granarius and Tenebroides mauritanicus are associated with cereals today.
However, the two beetles suggest different stable-carbon isotopic values for the cereals at Erkelenz187
Kückhoven. While the isotopic ratios may reflect trophic level differences, T. mauritanicus, as a
carnivore and an omnivore, should exhibit a more enriched carbon-13 ratio than the herbivorous S.
granarius. However, this was not the case in the present study. Unfortunately, δ2H and δ15N values
could not be procured for the cadelle specimen, due to limited fossil material, so interpretation is
limited. The δ13C ratio for Sitophilus granarius implies a higher altitude origin for the cereals (the
hinterlands to the north and west of the site are higher in elevation than Erkelenz-Kückhoven) whereas
the signature for Tenebroides mauritanicus is closely correlated with the species assumed to be from
the local environment. The discrepancy between the two grain associates has been interpreted in three
ways:
1) cereals were being cultivated near the village and in the higher altitudinal hinterlands;
2) cereal was being cultivated locally and being imported from a location higher in altitude or
lower in latitude;
3) cereal was being cultivated only in the hinterlands and represented by the granary weevil,
whereas the cadelle was not associated with the cereals.
The cereals were most likely being cultivated in different areas around the village; perhaps
different grains were grown at different altitudes. However, because of the variation between the two
grain associates, the possibility of cereal importation cannot be dismissed. The third suggestion
regarding Tenebroides mauritanicus not being regarded as a cereal associate is remote. While the
cadelle has been documented living beneath the bark in trees, those records are from warmer climates
(Crowson 1958; Palm 1959), and even though the species is considered cold hardy, it has not been
found in non-synanthropic habitats in the cold temperate regions (cf. Hunter et al. 1973) and exhibited
life cycle requirements inferring warmer climate origins [see Chapter 5].
7.4.8 Synopsis
This case study provides evidence of isotopic differences in insect species both within and
between sites in Linearbandkeramic Germany. The isotopic predictions indicated a highly variable
arthropod population, which was interpreted as evidence of the exploitation of the natural and
cultivated resources in the local environment and the surrounding hinterlands. Both the dung fauna
and plant-associates suggested that the domesticated vertebrate herbivores and agricultural plants were
tended away from the site. At Eythra, Plaußig, and Erkelenz-Kückhoven, the isotopic ratios associated
with cereals suggested that the agricultural crops were cultivated at higher altitudes than their
corresponding sites. Moreover, at Erkelenz-Kückhoven, the stable-isotope signatures of dung beetles
implied that the lower altitude regions may have served as grazing areas for the vertebrate herbivores.
While the isotopic discrepancies between the perceived local vegetation entomofauna and the grain
associates were interpreted as local versus hinterland exploitation, the enriched ratios evidenced by the
grain fauna may be indicative of the importation of cereals from lower latitude regions. The carbon188
13 variation reflected by grain-associates Sitophilus granarius and Tenebroides mauritanicus at
Erkelenz-Kückhoven may represent the importation of cereals in addition to the local ‘subsistence’
agriculture.
7.5 Conclusion
The stable-isotope analysis of insect remains has potential as a method for discerning human
activity in the past. Because the geographically distinct isotopic signatures of a region are assimilated
into insect chitin through diet and become locked into the chitin following adult metamorphosis,
recovered archaeological insect fossils are excellent isotopic indicators.
However, interpretation of the isotopic values requires an understanding of the ecology of the
insects. The trophic level variability of isotopes presented by vertebrate herbivores, omnivores, and
carnivores appears to be more convoluted in insects with the additional inclusion of mycetophagous
and, potentially, coprophilous species, which may further distort the isotopic signal, especially in
regards to carbon-13 and nitrogen-15 ratios. As interpretation of isotopic results is limited by an
understanding of species’ ecology, it is crucial to select species for analysis, which have habitats and
dietary habits that have been well documented.
The application of stable-isotopic carbon, hydrogen, and nitrogen on insects recovered from
West Stow and Neolithic Germany showed potential as a means of assessing local versus non-local
signatures. In the present study, low levels of isotopic fractionation were interpreted as differences in
seasonality or altitude, for which latter was construed as indicating a hinterland connection. In regards
to long-distance introductions, the Dorcus parallelopipedus specimen provided the only clear evidence
of a species of non-local origin although the Erkelenz-Kückhoven grain pests may be indicative of a
second case. While the lesser stag beetle most likely represents a natural invasive rather than a human
introduction, the species’ isotopic signature was distinct from the other species in the assemblage
clearly identifying it as non-indigenous. This suggests that isotopic analyses may be of use in
recognising species associated with long-distance exchangeable commodities in the past.
Unfortunately at this stage, the stable-isotopes recovered from archaeological insect remains
are unable to serve as a means of identifying the origins of foreign goods. While well-correlated, the
isotopic fractionation between the meteoric precipitation and the chitin is not back traceable to
geographic origins through linear models. Although Rubenstein and Hobson (2004) and Bowen et al.
(2005) offer non-linear interpolation methods which have proven successful in modern studies, the
models require an isotopic-geographic patterning correlated with the data set. Because insect remains
have not been as frequently analysed as vertebrates, the isotopic data necessary to back-trace or map
the species do not yet exist.
189
While the application of isotopic analysis towards insect remains is still in its infancy, it has its
advantages as a palaeoeconomic tool. Although the method operates on a similar premise as the
palaeoecological method discussed in Chapter 4, it advances beyond inferential assumptions of foreign
and local product-associates and provides tangible evidence towards the isotopic signal of the
geographic origin of the species’ (primary level) dietary source. The isotopic approach is also an
improvement to the biogeographical method as it by-passes the inherent limitations of a spatialtemporal approach [see Chapter 5]. Isotopic analysis offers inferences based upon the individual
specimens analysed and is not contingent upon the documented archaeological presence of the species
beyond the site.
190
Chapter 8
Recovery of DNA from Archaeological Insect Remains: First Results, Problems
and Potential
Modified from the Journal of Archaeological Science publication
Recovery of DNA from archaeological insect remains: First results, problems and potential.
Journal of Archaeological Science 36, 1179-1186 (2009)
Gary A. King a, M. Thomas P. Gilbert b, Eske Willerslev b, Matthew J. Collins c, d and
Harry Kenward a
a
Department of Archaeology, University of York, The Kings Manor, York YO1 7EP, UK
b
Biological Institute, University of Copenhagen, Universitetsparken 15, 2100 Copenhagen Ø,
Denmark
c
BioArch, Department of Archaeology, University of York, UK
d
BioArch, Department of Biology, University of York, UK
191
Abstract
We report the recovery of short fragments of PCR amplifiable ancient DNA from exoskeletal
fragments of the grain weevil Sitophilus granarius (L.) extracted from Roman and medieval deposits
in Northern England. If DNA preservation in archaeological insect remains is widespread then many
applications in the spheres of evolutionary studies and archaeology can be conceived, some of which
are outlined.
Key words: ancient DNA, thermal age, beetles, Roman, medieval, Sitophilus granarius
8.1 Introduction
The disarticulated remains of insects have long been successfully employed as bioindicators of
natural palaeoecosystems and in archaeological reconstructions (e.g. Coope et al. 1998; Buckland
2000; Coope 2000; Robinson 2001). The huge potential of archaeological insects as evidence of past
human activity and palaeoecology arises from their ecological diversity, their tendency to be ignored
or perceived as unimportant to humans, and their sensitivity and rapid reaction to environmental
change. The discipline of archaeoentomology works from the premise that in most temperate and
arctic environments of the northern hemisphere, all or most insects have maintained morphological
and physiological stability during the Quaternary (Coope 2004). On this basis, palaeoecological
information can be extracted by superimposing the climatic range and ecological role of modern
insects over the fossil record.
Here we report a preliminary investigation of the preservation of DNA in insect remains from
archaeological sites in Northern England. There are numerous reasons why analysis of DNA from
such fossils should be carried out, but three are perhaps the most significant. The first is to test the
hypothesis that morphological constancy can be equated with genetic, and thus presumably
physiological and behavioral, constancy. The second is to refine characterization and identification of
species, ‘races’, or other populations. While they are normally disarticulated, the preservation of
insect fossils is such that they typically retain morphological characteristics, including microsculpture,
and can be identified by comparison with modern reference material. However, the morphological
similarity of many closely related taxa limits identification to species group, genus, or even subfamily.
This is evident from inspection of most published species lists. In order to overcome the difficulties
associated with the classical methods, other means for identification must be investigated. Some
mainstream entomologists have been researching the benefits of genetic (DNA-based) identification
methods with a large degree of success; the method permits rapid and accurate determination,
regardless of developmental stage or specimen damage, and has even been proposed as a solution to
the problem of the shortage of skilled taxonomists (Tautz et al. 2003).
192
The third reason for analysing DNA from fossil insects is the possibility of recognising
intraspecific variations which may provide clues as to the past demography and history of dispersal,
whether natural or by humans, of species.
Where insects (and other organisms) have been
investigated, local populations have been found to have subtly different genetic characteristics. In
Europe, for example, these have been used to track postglacial recolonisation from glacial refugia
(Hewitt 1999, 2000, 2004). While problems have been signaled (Reiss 2006), recovery of DNA from
Pleistocene and archaeological insect fossils might thus open up significant new lines of research.
For the present investigation we have chosen the granary weevil, Sitophilus granarius. This is
an economically important beetle which is not a native of Britain, but whose remains are commonly
recovered from waterlogged archaeological deposits and have yielded amino acids in tests conducted
on Roman Age subfossils [Appendix 5].
A non-destructive method for ancient DNA (aDNA)
extraction (Gilbert et al. 2007) has been employed to demonstrate the presence of ancient DNA in
weevils from Roman and medieval deposits in Northern England.
8.2 Materials and Methods
Subsamples of raw sediment from 62-8 Low Petergate, York (excavated in 2004; Hall et al.
2007) and Park View School, Chester-le-Street, County Durham (excavated in 2006; Schmidl et al.
2006) were processed in 2007 using methods outlined by Kenward et al. (1980). Paraffin flotation
was employed for the extraction of insect remains, and the resulting ‘flots’ were stored in alcohol until
they were sorted using a low-power binocular microscope. Selected fossils were temporarily stored on
damp filter paper in closed dishes together with a few thymol (i.e. 6-isopropyl-m-cresol) crystals to
prevent mould, then placed in small glass vials containing industrial methylated spirit (an
approximately 90/10 mixture of ethanol/methanol). Fragments of S. granarius from six archaeological
samples were selected for DNA analysis, and modern specimens from a laboratory culture were used
as controls (Table 8.1). Precautions were followed to avoid contamination of the material with
previously amplified DNA (following Cooper and Poinar 2000, as modified by Gilbert et al. 2005).
Prior to Polymerase Chain Reaction (PCR) amplification, the DNA extractions and subsequent
manipulation were performed in a laboratory designated for research on samples containing low
concentrations of DNA, including aDNA. This laboratory is physically isolated from the laboratory
where post-PCR work is conducted. Furthermore, the PCR amplified DNA was both cloned and
sequenced from multiple overlapping PCR amplifications from each sample to ensure sequence
accuracy.
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Table 8.1 Details of specimens of Sitophilus granarius examined
Specimen
Fragments
Sample
Date
Amplifiable
Amplifiable nuDNA
mtDNA
GK1
Whole
Yorkshire
Modern
98bp
Y
specimen
GK5
57
Y
bp
14th c.
148bp
Y
88
256bp
Y
bp
98bp
Y
57
3 heads; 5
62-8 Low
elytra; 1 leg
Petergate, York
148bp
Y
bp
4977 76/t2
256
Y
88
Y
Y
Y
bp
GK6
8 elytra
62-8 Low
th
14 c.
98bp
Y
Petergate, York
2 elytra
Park View
Roman
148bp
Y
88
256bp
Y
bp
98bp
Y
57
School,
5 elytra
Park View
Roman
148bp
N
88
156bp
N
bp
98bp
Y
57
School,
1 head; 1
Park View
probiscus; 4
School,
legs
Durham 152
N
N
N
bp
Durham 144
GK10
Y
bp
Durham 113
GK8
Y
bp
4977 76/t1
GK7
57
Roman
148bp
N
88
256bp
N
bp
98bp
N
57
N
N
bp
148bp
N
88
256bp
N
bp
N
DNA is expected to degrade rapidly following cell death, limiting the potential PCR
amplifiable size of extracted DNA template molecules (Lindahl 1993a). Therefore to investigate the
relative preservation of mitochondrial DNA (mtDNA) and nuclear DNA (nuDNA) in the
archaeological samples, the PCR assays were designed to screen for DNA survival at a range of
fragment sizes in order to assess thermal age [see Figure 8.1]. For detailed methods and arguments for
data authenticity see Appendix 6A.
194
Figure 8.1 Comparison
mparison of amplification success of different amplicons when converted to thermal age. Results are
shown for mtDNA (ν
ν), and for nuDNA where the ratio of target numbers to mtDNA is either 1:100 (λ
(λ) or 1:2 (
)
8.2.1 Thermal age: Accounting for Target Length and Copy Number
Thermal age (Smith et al. 2003) attempts to normalise different samples for the effects of
burial temperature and time. Thermal age was estimated aassuming
ssuming an activation energy of (127.8 kJ
mol-1) giving an approximate rate for DNA depurination at 10 °C of 4 x 10-6 yr-1.
Thermal age is a useful way of contrasting the relative likelihood of amplification success of
different initial copy number (sample type, target) and different amplicon lengths. The number of
sites of potential depurination, which increases with amplicon length, increases the probability of
chain scission (Deagle et al.. 2006). By normalising to an amplicon length of 100 bp (including
primers), samples from the same site and of the same age will have different thermal ages that reflect
target size (and hence
nce probability of depurination).
The number of targets surviving is also a function of starting copy number. If one tissue type
has more target copies than another (e.g. muscle versus bone), or if mitochondrial and nuclear targets
are analysed from the same
ame sample, these targets will have different thermal ages (assuming that rates
of chain scission do not differ). For example nuDNA of equal length to mtDNA will have a higher
thermal age, because the lower initial starting copy number implies fewer targe
targets
ts will be present in the
sample for any given time interval. In the absence of estimates of absolute concentration in recently
buried insect cuticle, the differing relative concentration of targets such as mtDNA and nuDNA can be
explored. Equivalent thermal
rmal ages imply equal probabilities of successful amplification.
8.3 Results
Mitochondrial DNA was successfully extracted, PCR amplified, cloned and sequenced from
five of the six specimens. While the modern and medieval samples yielded mtDNA at all the fragment
195
lengths investigated, only the shorter 98 bp mtDNA fragments could be amplified from two of the
three Roman specimens, with the last yielding no amplicon. Similarly, while short nuDNA fragments
were successfully extracted and sequenced from the medieval and modern samples, nuDNA
preservation was not identified in the Roman fossils. No PCR or extraction blanks exhibited evidence
of contamination. The identity of the sequenced DNA as Sitophilus granarius was confirmed both
through the modern control sample sequences, and through positive identification against the NCBI
GenBank database. Thermal age estimates were very similar for samples from York and Durham.
The former could be compared with a direct measurement from measured soil temperature (at 4 m
depth, 10.9 °C; Chang 1958) giving an estimate of 1.17 times the rate at 10 °C.
Normalising for fragment length, data from mtDNA amplifications suggests that the thermal
limit (the point at which no amplification is successful) for mtDNA is an order of magnitude lower
than estimated for bone. The small sample size will reduce the errors on this estimate, but would not
account for such a large difference in thermal limit. Assuming a very conservative estimate of the
ratio of mitochondrial to nuclear DNA (1:100), thermal age estimates for nuDNA are an order of
magnitude higher than mtDNA. It is necessary to reduce the ratio to 1:2 in order that the thermal
limits overlap. If we discount the possibility that this ratio is an inaccurate estimate of starting copy
number, then the result indicates that the respective success rates of mtDNA and nuDNA does not
correspond to an equivalent chemical (i.e. temperature/time dependent) process.
The two most
probable explanations for this are (i) differential rates of depurination due to differences in the packing
of the two types of DNA within insect remains or (ii) additional biological (nuclease activity) which
has preferentially targeted nuDNA. The results do suggest that nuDNA is preferentially preserved
relative to mtDNA in these samples.
8.4 Discussion
We have shown that PCR amplifiable levels of mtDNA and multicopy nuDNA survive in the
exoskeletons of fossil beetles. Preserved insect DNA has been reported from amber-encased fossils
dating to the Ogliocene (25-35 mya), e.g. stingless bees (Cano et al. 1992a, 1992b), termites (DeSalle
et al. 1992, 1993), and wood gnats (DeSalle and Grimaldi 1994), and the Cretaceous (120-135 mya),
e.g. Lebanese weevils (Cano et al. 1993).
However, as few recent studies have succeeded in
amplifying DNA from remains older than several hundred thousand years, it is suggested that the
results of amber studies should be regarded carefully.
Investigations of sub-glacial deposits in
Greenland (Willerslev et al. 2007) and insect carapaces from museum collection samples (less than
one hundred years old, e.g. Zakharov et al. 2000; Junquiera et al. 2002; Gilbert et al. 2007) have
yielded DNA fragments with no indication of contamination. While previous analyses have been
conducted with varying degrees of success, the recovery from terrestrial fossils in archaeological
196
deposits opens up many new possibilities, providing preservation is fairly common and occurs in a
range of other taxa.
Intraspecific variation of DNA between modern local populations and races has been
demonstrated for a number of insect taxa, including honey bees (e.g. Garnery et al. 1995; Arias and
Sheppard 1996; Lee and Hall 1996), ground beetles (e.g. Ashworth 1996; Reiss et al. 1999; Cardoso
and Vogler 2005), some other groups of beetles (e.g. Schrey et al. 2005; Smith and Farrell 2005),
human lice (Leo et al. 2002; Yong et al. 2003), and butterflies (e.g. Nice et al. 2005). All of these
studies of modern genomes have been relevant to palaeoecological questions, but all lack an essential
element: information about the genetic constitution of ancient populations. Given the success of
genetic analysis of modern insects, its application to subfossil insect remains clearly deserves
exploration.
In the present study, ancient DNA was successfully extracted from one species of beetle from
Roman and medieval contexts. The failure to amplify nuDNA and longer strands of mtDNA from the
Roman material is consistent with what would be expected as a result of postmortem fragmentation of
the DNA (Lindahl 1993a; Deagle et al. 2006). However thermal age analysis suggests that the thermal
age limit at which no amplification success is possible (for a nominal 100 base pairs at a constant 10
°C) is at least an order of magnitude less for DNA in these insect remains (2-3 ka) than in bone (Smith
et al. 2003). This analysis also reveals that nuDNA is relatively more resistant to hydrolysis than
mtDNA. The reasons for these two observations are as yet unknown, but together they do imply a
significantly greater role of biological processes (nuclease) activity in the destruction of DNA in these
remains when contrasted with available data from bone (e.g. Poinar et al. 2006). The reason that
specimen GK10 did not yield mtDNA of 98 bp lengths while other material of similar age gave good
recovery is unknown, but given probable role played by nuclease activity in polymer scission, this
may reflect early taphonomic processes. However, the GK10 fossils did not demonstrate any visible
signs of morphological degradation which differed from specimens GK7 and GK8.
If DNA can be recovered from S. granarius, it is likely to be preserved in many other insects,
at least in those with similarly substantial exoskeletons. A priority is therefore to test a wider range of
robust fossils, and to investigate the possibility that DNA may survive in more delicate fossils as well.
For the mtDNA analysis used here, we targeted a conserved portion of the cytochrome oxidase I gene
for two predominant reasons. Firstly, it has previously been sequenced in modern samples, so that a
reference sequence was available against which to design primers and compare the data (O’Meara and
Farrell unpublished; Genbank ID: AY131101); and, secondly, its relatively conserved state within
species suggested that the ancient sequences were unlikely to be sufficiently different from the modern
sequences such as to lead to PCR amplification problems. However, the conserved nature of that gene
segment also means that its use for characterizing populations or tracing micro-evolutionary change is
likely to be limited. Therefore, for future studies the analysis of more variable genes or at least a more
197
variable portion of the cytochrome oxidase I gene (see Juan et al. 1998) will need to be employed.
One potential candidate is the mtDNA cytochrome oxidase II gene, which has recently been used by
Moya et al. (2004) in phylogenetic investigation of, and discrimination between, ground beetles of the
genus Eutrichopus. Additionally, Juan and collaborators (1998) have had success utilizing the 255
base pair fragment of the cytochrome oxidase I gene that coincides with the 5’ end positions of the
2410 and 2665 in the Drosophila yakuba mitochondrial genome to formulate the phylogeography of
the darkling beetle Hegeter politus in the Canary islands.
How might we use DNA from ancient insects?
Numerous taxonomic, evolutionary and
archaeological applications spring to mind. It may prove useful for crucial identifications, although
there are probably few cases where the effort would be justifiable; one exception might be the
differentiation of races of human lice (see below). The possibility of detecting minor genetic change
through time (microevolution), whether gradual, or perhaps in sudden steps, such as when new
selection pressures were applied when populations were translocated geographically or into artificial
environments, would open up the opportunity to make comparisons with genetic changes seen
following invasions at the present day (e.g. Huey et al. 2000).
The morphological constancy of
insects through the Quaternary is ascribed to remixing of genetically divergent populations by climate
change. Although some PCR amplifiable insect DNA may be expected to survive in deep frozen (i.e.
permafrost environments), routine recovery of Pleistocene insect DNA is perhaps too much to hope
for. However if it were achieved, for example using fossils from tundra deposits, we should be able to
trace this process of differentiation and rehomogenization through time.
Do modern natural and synanthropic (human-associated) insect populations differ genetically,
can this be seen in ancient populations, and if so, does this indicate adaptation or founder effect in
isolated populations (cf. Frankham et al. 1999)? More fundamentally, do the genomes of fossils differ
substantially from modern examples, and if so can we reasonably continue to assume that past
populations exploited similar habitats to present ones and can thus be used in reconstructing past
ecology? Specific problems might be soluble: for example, the ground beetle Pterostichus madidus
(Fabricius) is very common today, but rather rarely found as a fossil: why did it become more
common? Was there a genetic change (i.e. adaptation) associated with increasing synanthropy? There
are some ‘sibling species’ among insects which appear to have identical habitats and which often
occur together. A case is provided by Anthicus floralis (Linnaeus) and Anthicus formicarius (Goeze),
found in materials such as stored hay and both with a substantial fossil record. Might the origin of
such pairs be traced, and the cause (adaptation or isolation?) be determined? A third case worthy of
note is the head and body races of human lice, whose origin seems on the basis of studies of the
modern genomes to be complex, the races perhaps having arisen twice independently (Leo et al. 2002;
Yong et al. 2003); might fossils clarify the relationship between them? Perhaps one of the most
valuable uses of genetic analysis of fossils in archaeology might be in characterizing past local
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populations and tracing patterns of natural or human distribution. The sources of imported materials
such as dye plants might conceivably be traced through recognition of geographically localized
genotypes of insects associated with natural-habitats. The history of synanthropic pests such as S.
granarius, itself, might be traceable: for example, were British populations derived from a small
number of Roman introductions, or established and maintained by continued trade?
Did the
population present following the Norman Conquest arise from British survivors of the ‘Dark Ages’, or
from a new introduction from Continental Europe?
Single or multiple introductions to a settlement might stand as evidence of intensity of trade (a
parallel to work on synanthropic insects, Kenward 1997). To cite a specific case, an Early Christian
site in Country Antrim, Northern Ireland, seems to have had far more synanthropes than would be
predicted from its apparent isolation (Kenward 1997; Kenward et al. 2000). Was this a result of
intense exchange over a period of time (multiple genotypes being predicted), or of one large-scale
introduction of small numbers of a range of synanthropes, in hay for example (restricted range of
genotypes predicted)? As with studies of DNA generally, it is suspected that, providing preservation
is widespread, many new uses will arise.
8.5 Conclusion
In this study the investigation of mtDNA and multicopy 18s nuDNA survival from the anoxic
waterlogged remains was conducted on the granary weevil Sitophilus granarius as part of a pilot study
for on-going research. The genes were selected based on the existence of GenBank records and their
genetic sustainability. As previously noted, the portion of the CO1 gene was selected for analysis
because of its conservative nature and consequentially reduced chance of variability due to population
and specific mutations overtime. 18s was selected because it is found in multiple copies, not to the
extent of mtDNA, but more than the single copy of other nuDNA genes. It is likely that future studies
will to some extent benefit from also testing whether single copy nuDNA survives in similar samples,
although this is likely to be sample dependent, as indicated by the variation in the medieval and
Roman material recorded here. We believe that the results conveyed in the present study are authentic
and that the successful recovery of ancient DNA from waterlogged insect remains indicates significant
potential.
8.6 Acknowledgements
We are grateful to Terry O’Connor and Ellie Jones for their comments on a draft. A special
thanks to PCA North and Palaeoecological Research Services for supplying Roman samples from the
Park View School site. GAK’s research was supported by the Short-term Marie Curie PALAEO
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Fellowship and the kind contributions from the Department of Ancient DNA and Evolution at
University of Copenhagen. HK was co funded by English Heritage and the University of York.
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Chapter 9
Conclusions and Future Directions
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9.1 Summary
The field of palaeoentomology is rich in unfulfilled potential and promise. The study of insect
remains retrieved from archaeological contexts has been particularly effective in reconstructing past
ecologies and living conditions. Furthermore, it presents the opportunity to investigate an exciting
range of previously unaddressed or un-approachable questions concerning past social and economic
activities; an endeavour which was explored in the present study.
The effectiveness of palaeoentomology as an interpretative tool relies on an understanding of
insect species’ morphology and physiology.
Chapter 2 provides a brief survey on the role of
morphological features in species identification and behavioural interpretations. Additionally, an
understanding of cuticular composition and formation is invaluable in the successful design and
performance of biochemical and biomolecular extractions [Chapters 6, 7, and 8]. In order to discern
information regarding past human activity, it is essential to understand how an organism interacts with
its environment, its trophic position, and ecological constraints [Chapter 2].
In an effort to explore the potential of palaeoentomology as a palaeoeconomic tool, three
analytical approaches were tested—palaeoecology, biogeography, and isotopic analyses—and a fourth
a method was proposed, i.e. phylogeography.
Each of the tools had distinct advantages and
limitations.
9.1.1 The Palaeoecological Approach
Through analysis of species’ morphology and ecological preferences, palaeoecological
reconstructions may be formulated and potential socio-economic indicator species pinpointed [Chapter
4]. The palaeoecological approach to discerning past trade and human movement involved:
1.) The processing of environmental samples and the identification of insect remains from a single
archaeological site;
2.) The interpretation of the retrieved arthropod fossils to discern the habitat preferences and diet
of individual species. The information was employed to compile palaeoenvironmental
reconstructions, which aided in interpreting the environment at and near the site.
The
abundance of insect remains and species ecological groups served to identify autochthonous
and allochthonous components;
3.) The evaluation of the preferred habitats and diets of modern species to draw parallels with the
archaeological fauna.
The ecological preferences of the modern taxa were viewed as
analogous and thus were superimposed over the archaeological specimens in order to identify
insect species associated with human exploitable commodities;
4.) The application of the Mutual Climatic Range model to calculate the temperature range at the
site for the warmest and coldest months. The results of the model were based on the temperate
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requirements and geographic range of carnivorous and scavenging beetles. It is important to
note that in the present study, the insect species used as indicators of human activity were
primarily stenotopic herbivores and/or ectoparasites. Thus the selected indicator fauna was
independent of the fauna used in the MCR model, which means the temperature predictions of
the model were not influenced by potentially foreign species; and,
5.) The comparison of the determined palaeoenvironmental and palaeoclimatic signature of the
site with the geographic and thermal requirements of the proposed indicator species. These
evaluations were used to hypothesise as to whether the species could have inhabited and
survived in the ambient environment.
The palaeoecological approach provides a comparatively quick and non-destructive means of
identifying insect species that may stand as direct or indirect evidence of human-driven socioeconomic activities, including probable evidence towards the exploitation of local versus foreign
commodities. The method suffers in that it is highly speculative and can be unconvincing in its
conclusions.
While it can effectively target taxa associated with exploitable commodities, e.g.
honeybees implying the availability of honey and beeswax, a single site palaeoecological assessment
lacks the ability to provide a timeframe for the introduction of the species.
For example, the
Damalinia ovis recovered from 16-22 Coppergate was taken as a strong indication of the presence of
wool in Anglo-Scandinavian York. Based on its thermal requirements, it was identified as a species
that was not indigenous to the United Kingdom and which was adapted to a warmer climate.
However, does this imply that the Vikings were importing sheep and/or wool from warmer geographic
regions, or was the species introduced to Britain during an earlier period and able to endure in microclimates? The palaeoecological approach cannot account for the latter possibility. Furthermore, how
would palaeoecological approach rationalise the predominantly northern fossil record of the sheep
louse (e.g. Perry et al. 1985; Buckland et al. 1992; Kenward and Allison 1994a; Schelvis and Koot
1995; Buckland et al. 1998)?
The other major limitation of the palaeoecological method, as utilised here, is its difficulty in
ascertaining the origin of the exchangeable resources. Species, such as Apion (Exapion) difficile,
which were capable of surviving in Britain and have a modern British presence, are assumed to be
representative of the local exploitation of resources. However, the Anglo-Scandinavians, in the case
of A. difficile, may have introduced the weevil with imported dye plants from Continental Europe,
where it may have been much more abundant. Fortunately, the trade signal is not always completely
ambiguous when using this approach. For example, in the case of the Hesperophanes fasciculatus
recovered from Roman Alcester (Osborne 1971), the species, which is associated with structural
timbers and furniture and believed unable to survive in Britain, has a modern distribution extending
through southern Europe and the Middle East, and is particular common in Greece. The thirteen
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individuals of H. fasciculatus in Roman Britain are indicative of maritime connections with the
Mediterranean.
Despite the method’s limitations, the palaeoecological assessment has two key advantages as
tool for indicating trade and cultural contact.
1.) It is relatively cost-effective. The approach does not burden the researcher with excessive
laboratory fees. It requires access to a water source, a bucket, sieve, and paraffin, and some
costs may be incurred if travel is necessary for specimen identification or document retrieval;
and
2.) It also produces fairly timely results. Single sample processing can usually be achieved over
the course of a few hours, but can vary depending on soil matrix and weight. The time
involved in specimen identification and palaeoecological reconstruction is relative to the
experience of the analyst and the number of fragments present in the flot.
The outcome of both factors may vary between sites and samples. However, they constitute
the core aspects of the palaeoecological approach. Furthermore, the other analytical methods must
also account for these factors, especially the processing and identification costs, as they comprise part
of each of the methodological approachs. Thus, the palaeoecological approach provides the quickest
and most affordable results.
9.1.2 The Biogeographical Approach
Through employment of the biogeographical method, a few of the limitations of the
palaeoecological approach may be addressed.
Palaeobiogeographical investigations rely on
archaeological, ecological, and historical accounts to map changes in a species’ distribution through
time and space [Chapter 5].
This enables the researcher to discern a timeframe for the initial
introduction of species to a region. The biogeographical approach involved the following steps:
1.) The selection of indicator species. In the present study, grain-associates were used;
2.) The compilation of historic and archaeological data on the species. These were used to
evaluate the location of each beetle at different points in time;
3.) The examination of the temperature specifications of each species. Because most members of
the grain fauna have a cosmopolitan distribution today, the thermal conditions necessary for a
species to complete the development of its life cycle were viewed as an indication of the
probable temperature range of its native region, i.e. the area where it evolved and was adapted
to survive in the wild; and,
4.) The archaeological, ecological, and historical information was correlated to determine when a
grain species first entered a particular geographic region and whether the species may have
been endemic to that area.
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By applying the biogeographical method to the grain pest fauna, it was possible to glean an
understanding of human activity in the past. Though there are exceptions, the grain beetles, in general,
tend to be both strongly synathropic and poor dispersers. For example, Sitophilus granarius only
infests cereals that have been harvested and stored. Additionally, S. granarius does not fly, which
would impede its effectiveness as a disperser, especially across geographic barriers, e.g. bodies of
water, deserts, mountains, etc. Although the weevil may have been able to self-disperse between
habitats via corridor pathways, filter and sweepstake routes would have presented obstacles. Thus the
species wide-spread distribution, even as early as the Neolithic, is likely to have been attributed to
anthropogenic methods, i.e. the species would have hitchhiked to various locations in cereals carried
by man.
Because of this anthropogenic connection, the biogeographical interpretation of the
distribution of the grain-associated insect fauna may effectively stand as evidence of human
movement through trade, culture contact, or migration.
As mentioned in Chapter 5, the major limitations of the biogeographical approach are:
1.) It is unable to distinguish between multiple introductions of the same species to a region and,
as such, is only able to imply contact between cultures or regions that involved the ‘initial’
introduction of the pest; and,
2.) Its effectiveness is restricted to the accessibility and availability of reliable archaeological and
historical accounts.
Although the method has some limitations, it is a proficient means of discerning human
movement in the past. By examining the geographic distribution of an indicator species at a set period
of time, it is possible to propose likely points of origin for trade or culture contact. For example, the
hypothesis that the grain from the late second century ship, which was excavated in the Netherlands
(Pals and Hakbijl 1992), originated in Britain based upon the grain fauna ‘community’ that was
present in the assemblage [Chapter 5]. This is a distinct improvement over the palaeoecological
method which would only have been able to recognise the grain pests as potential allochthonous
species and suggest a connection to warmer climates on the basis of their thermal requirements.
The biogeographical approach is also fairly cost effective.
Although it may involve the
processing of environmental samples, the method is primarily dependent upon the review of available
literature. While this may be time consuming depending on the species being assessed, it is a
relatively low cost means of exploring potential trade connections and culture contact as made evident
through insect subfossils.
9.1.3 The Isotopic Approach
The isotopic analysis of insect remains is not a well-established field of research. Although
isotopic studies (e.g. Miller et al. 1988; Gröcke et al. 2006) have proposed the application of insect
fossils to palaeoclimatological research, few attempts have been made (e.g. Wooller et al. 2004;
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Hardenbroek 2006; Hardenbroek et al. 2007). Despite this, isotopes extracted from modern insects
appear to be well-correlated to the meteoric precipitation of their environment, especially stable
carbon and hydrogen (cf. Miller 1984; Hobson et al. 1999), which suggests their potential as
palaeoenvironmental indicators. Furthermore, Chapter 6 demonstrated the ability of insects, under
laboratory conditions, to assimilate and retain the isotopic signature of their host plant’s (comprising
both foreign and locally grown cereal crops) region of origin, which implies the ability of insect
remains to stand as evidence of past trade. In Chapter 7, stable-isotopic carbon, hydrogen, and
nitrogen were examined to pioneer the application of isotopes from archaeologically recovered insect
remains toward discerning palaeoeconomic activities. The isotopic approach involved:
1.) The recovery of insect remains from modern and Neolithic sites;
2.) The isolation of the chitin to avoid the potential of assaying conflicting isotopic signatures
from proteins and lipids (see Miller 1984). This is significant as it involves the destruction of
the insect fossil, which needs to be taken into consideration when applying the method to
specimens, rare or otherwise;
3.) The extraction of stable isotopes from the insect chitin. Carbon-13, deuterium, and nitrogen15 were assayed; and,
4.) The comparison of the isotopic signatures of species within and between sites; accounting for
trophic level variation.
The isotopic method proved an effective means of assessing past socio-economic activities.
The analyses denoted isotopic variation within the modern and Neolithic sites. In most cases, the
isotopic signature procured from the cereal- associated species differed from the ratios of the
entomofauna that were presumably feeding on materials endemic to the site. The discrepancy was low
and was interpreted as expressing exploitation of resources from the local hinterland. The isotopic
variation was most likely attributed to differences in altitude or seasonality. Although the present
study did not recover an indication of a long-distance economic connection from the archaeological
sites, the stable-isotopes extracted from Dorcus parallelopipedus, which based on the isotopic results
was assumed to be a natural invasive to the Neolithic site, evidenced the ability of the method to
distinguish between local and potentially foreign signatures. From the modern site of West Stow, the
isotopic ratios from Anobium punctatum and Sitophilus granarius may imply importation of building
materials to the site, i.e. wood for structural timbers and cereal for thatch, respectively. However, the
discrepancy between the autochthonous and allochthonous species from West Stow was not sufficient
to assume the foreign importation of the products, rather it merely suggested that the structural timbers
and thatch materials originated beyond the site.
Because of the paucity of previous isotopic analyses on archaeoentomological materials, the
isotopic method suffers from an inability to back trace the origins of potentially foreign materials.
Thus while the approach can provide convincing, if not conclusive, evidence as to whether a
206
commodity was local or imported, which is an improvement over the speculations of the
palaeoecological approach, it is unable to identify a port of origin. If more archaeoentomological
evidence was assayed in the future, non-linear interpolation methods based on geographic-patterning
(e.g. Rubenstein and Hobson 2004; Bowen et al. 2005) may be employed to back trace the materials.
The other major disadvantage of the isotopic method over the palaeoecological and
biogeographical approaches is cost. In addition to the minor expenditure involved in the processing of
the environmental samples, the isotopic assays incur costs (sometimes in the range of thousands or
tens of thousands of pounds sterling) from the purchase of chemicals and tin and silver capsules as
well as the fees required for the use of the laboratory equipment necessary for analysis and
computation. Depending on where the analyses are conducted, the isotopic method may require the
assistance of grants or other sources of funding. Although potentially costly, the isotopic approach has
potential as tool for discerning the localised exploitation of resources, trade, migration, and culture
contact in the past.
9.1.4 The Phylogeographic Approach
Chapter 8 presents the seminal discovery of ancient DNA from insect fossils recovered from
waterlogged archaeological contexts and outlines myriad lines of research that would benefit from its
application.
Unfortunately, a full phylogeographic analysis of archaeologically retrieved insect
specimens was not within the scope of this work.
However, previous studies have shown the
applicability of modern insect remains towards discerning phylogeographic relationships (e.g. Juan et
al. 1998).
As a tool for discerning the introduction of indicator species in the past and thus culture
contact, it is hypothesised that species derived from different populations will yield variation within
their genetic code. In the modern darkling beetle, Juan and associates (1998) determined a variation to
be present with in the CO1 gene. Moya et al. (2004) observed mitochondrial variation within the CO2
gene of the ground beetle genus Eutrichopus. If genetic discrepancy is evidenced in species recovered
from the same archaeological context or sample, it may indicate the introduction of the species to the
site, and thus may arguably stand as evidence of trade or culture contact, especially if the species is
associated with a resource which is exploitable by man.
In order to effectively back trace the origins of a species, phylogeographic methods need to be
employed. This requires knowledge of the region of genetic variability from populations of the same
species that are established in different locations. Ideally, this information would be derived from
contemporaneous populations.
For example, a project wishing to determine the origin of the Sitophilus granarius specimens
from 21 St. Peters Street Colchester, UK (King and Hall 2008), would attempt to compare the genetic
code from the Roman Colchester samples to the Roman Age specimens recovered from 1 Poultry,
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London, UK (Rowsome 2000), Santa Pola, Spain (Moret and Martin Cantarino 1996), Alphen aan den
Rijn, Netherlands (Kuijper and Turner 1992), Neuss, Germany (Cymorek and Koch 1969),
Touffréville Calvados, France (Ponel et al. 2000), Herculaneum, Naples, Italy (Dal Monte 1956), and
potentially the Iron Age specimens from Okruglo, Croatia (Smith et al. 2006) and Horbat Rosh Zayit,
Israel (Kislev and Melamed 2000). By comparing the S. granarius recovered from the different sites,
the distinct genetic signature of each population could be ascertained. If the Colchester and London
populations were identical, then it could be assumed that S. granarius was, at least originally, probably
introduced with cereals derived from a single geographic point. However, if the populations differed,
it may imply that Romans were importing cereals from different locations. By analysing the European
and Mediterranean populations, it may be possible to infer the source population and thus the origin of
both the Roman Colchester S. granarius and likely the cereals with which it was introduced.
However, caution would need to be exercised in interpreting data as the Sitophilus granarius from the
other regions may also have been imported [see Chapter 5].
While untested, the phylogeographic approach has potential as tool for assessing social and
economic activities in the past. As with the isotopic analyses, the method can be relatively expensive,
particularly in comparison to the biogeographical and palaeoecological methods. However, it is nondestructive method which may be capable of providing ‘definitive’ answers to otherwise unapproachable questions.
9.2 Future Directions
Various improvements to the methods and research approaches would enable significant
progress to be made as to effectiveness of palaeoentomology as research tool for assessing past trade,
migration, and culture contact.
Underpinning the success of future endeavours is the need for
increased palaeoentomological research in geographic areas which have received little or no prior
attention.
For example, the present study was severely impeded by the absence of
archaeoentomological material from regions such as North Africa and India, and suffered due to the
paucity of materials from continental Europe, the Middle East, and the Far East. The lack of subfossil
insect studies results in gaps in the palaeoentomological record, which in turn, limits the effectiveness
of the methods in being able to discern geographic and ecological patterns.
The application of ancient genetics and isotopes towards the study of archaeological insect
remains are nascent fields of research. As it has now been proven possible to extract uncontaminated
aDNA from waterlogged preserved insect fossils, concerted efforts need to be made to test the
applicability of a phylogeographic method towards addressing palaeoeconomic issues. Modern taxa
should be analysed to determine regions of intraspecific genetic variation, and further ancient DNA
should be extracted from archaeological specimens to determine geographic patterning within species.
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Can ancient DNA be recovered from pre-Roman waterlogged specimens? In addition, there is a need
to evaluate the survival of aDNA in charred and desiccated insect remains as they comprise a fair
proportion of the archaeoentomological remains from some regions. Moreover, the isotopic method
would be greatly advantaged by investigation of archaeoentomological remains from additional
archaeological sites. The procurement of further isotopic data would enhance the approaches ability to
back trace the origins of allochthonous species.
It may also be a worthwhile endeavour to apply holistically the methods addressed in this
investigation to more recent archaeological sites. Such an assessment would likely foster a clearer
understanding of migration, distribution, and origins of production through the multifaceted
establishment of spatial and chronological patterning. Consequently, contextual patterning may be
evident through the incorporation of evidence from associated historical and archaeological arenas.
The Colonial Period settlements in the Americas have recently proven rich in archaeoentomological
remains (e.g. Bain 1997, 1998; Bain et al. 2009; King et al. 2010). In assessments of the modern
fauna, Sailer (1983) estimated the presence of 1683 immigrant arthropod species in the continental
United States, of which 66 % are believed to have originated in the Palaearctic eco-zone, and Lindroth
(1957) claimed that 14 % of the Newfoundland ground beetles were European.
As the
archaeoentomological remains from Colonial Era sites are more recent than those evaluated in King et
al. (2009), there is the potential for both mitochondrial and nuclear DNA survival as the basepair
sequences would be expected to have undergone comparatively less decay than in the Roman and 14th
century specimens. The presence of longer, less fragmented DNA sequences would enhance the
likelihood of finding key regions of intraspecific variability which would be invaluable to
phylogeographic analyses. While individually each approach has proven informative in its own right,
a holistic assessment integrating the detailed results and analyses of all the methods should be
considered.
9.3 Conclusion
The methods reviewed in this text explored the potential of insect remains to stand as evidence
of past human interaction as well as to discern the origins of potentially foreign commodities. It is
suggested that through scrutiny of their ecological preferences, archaeologically retrieved insect
species may be relied upon to represent the presence of human exploitable resources.
Three
methodologies were tested to back trace the origins of the indicator species. The approaches differed
in the degree of confidence with which they could identify autochthonous and allochthonous species
and propose potential origins.
Additionally, a fourth approach was suggested but not applied.
Although the methods have limitations, it is proposed that palaeoentomological remains can be used as
a means of assessing culture contact. Furthermore, their effectiveness will be improved as future
209
studies expand the availability of applicable data sets, which will enhance the resolution of the
methods by enabling comparative geographic patterning.
210
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Appendices
250
Appendix 1
Mutual Climatic Range Data
251
Appendix 1A:
Species
TmaxHi
TMaxLo
TminHi
TminLo
TRangeHi
TrangeLo
MCR Species from 7-15 Spurriergate York
Dyschirius globosus
(Hbst.)
31
9
16
-47
57
8
Trechus obtusus/
quadristriatus (Er.)/
(Schr.)
36
10
31
-11
30
5
Pterostichus melanarius
(Ill.)
26
10
11
-37
49
7
Helophorus aquaticus/
grandis (L.)/ (Ill.)
36
10
31
-20
36
5
Cercyon tristis (Ill.)
24
9
9
-44
53
9
Megasternum obscurum
(Marsham)
26
9
13
-17
30
7
Xylodromus concinnus
(Marsham)
29
10
17
-16
30
7
Anotylus rugosus (F.)
28
11
10
-23
39
8
252
Anotylus sculpturatus
(Grav.)
30
15
17
-13
32
8
Anotylus nitidulus
(Grav.)
33
10
17
-43
53
8
Platystethus arenarius
(Geoff.)
18
9
7
-47
56
8
Platystethus nitens
(Sahl.)
33
15
15
-33
48
8
Leptacinus pusillus
(Steph.)
27
12
17
-27
44
9
Tachinus laticollis
(Grav.)
27
9
14
-35
44
7
Aphodius fimetarius
(L.)
29
9
15
-40
54
8
(formulated using Buckland and Buckland 2006)
253
Appendix 1B:
Environmental
Report
Overlap
NSPEC
TrangeHi
TRangeLo
TMinHi
TMinLo
TMaxHi
TMaxLo
MCR for 3 Roman Age Sites
Site
Kenward et al.
1986
Bedern Well
16
18
-8
6
11
24
37 97.30
Osborne 1971
Alcester,
Warwickshire
15
18
-7
6
11
24
15 93.33
Allison and
Kenward 1987
Copthall Ave.,
London
15
18
-7
6
11
22
30
100
(adapted from Table 1, King 2008)
254
Appendix 1C:
TrangeLo
TRangeHi
TminLo
Species
TminHi
TmaxHi
TMaxLo
MCR Period 4B Tenement C16-22 Coppergate Samples
Clivina fossor (L.)
29
9
18
-42
54
11
Trechoblemus micros
(Hbst.)
25
13
10
-26
43
10
Bembidion gilvipes
(Sturm)
23
15
9
-34
49
11
Bembidion biguttatum
(F.)
29
15
16
-18
33
10
Harpalus rubripes
(Duft.)
29
14
13
-19
37
9
Pterostichus melanarius
(Ill.)
26
10
11
-37
49
7
Megasternum obscurum
(Marsham)
26
9
13
-17
30
7
255
Hydrobius fuscipes (L.)
26
8
13
-53
62
8
Omalium rivulare
(Payk.)
28
9
18
-21
30
7
Xylodromus concinnus
(Marsham)
29
10
17
-16
30
7
Anotylus rugosus (F.)
28
11
10
-23
39
8
Anotylus sculpturatus
(Grav.)
30
15
17
-13
32
8
Anotylus nitidulus
(Grav.)
33
10
17
-43
53
8
Platystethus arenarius
(Geoff.)
18
9
7
-47
56
8
Platystethus nitens
(Sahl.)
33
15
15
-33
48
8
Leptacinus pusillus
(Steph.)
27
12
17
-27
44
9
256
Gyrohypnus
fracticornis (Müll.)
27
11
15
-17
30
8
Tachinus rufipes (L.)
27
9
15
-27
36
7
Aphodius prodromus
(Brahm)
29
12
14
-23
41
8
(calculated using Buckland and Buckland 2006)
257
Appendix 1D:
Environmental
Report
Overlap
NSPEC
TRangeHi
TRangeLo
TMinHi
TMinLo
TMaxHi
TMaxLo
MCR data from three Neolithic Age sites in Germany
Site
Schmidt 1998,
2010b
ErkelenzKückhoven
15
25
-11
9
10
28
8
100
Plaußig
16
24
-11
9
11
27
11
100
Schmidt 2010a
Schmidt 2005
Eythra
17
24
-10
9
13
30
7
100
(determined using Buckland and Buckland 2006)
258
Appendix 2
Evaluation of Biological Remains from a Roman Timber Drain at 21 St Peters
Street, Colchester (site code: 2007.124)
Modified from the Reports from the Centre for Human Palaeocology publication
Evaluation of biological remains from a Roman timber drain at 21 St Peters Street, Colchester
(site code: 2007.124).
Reports from the Centre for Human Palaeoecology, University of York 2008/15. (2008).
Gary A. King and Allan Hall
259
Appendix 2A:
Evaluation of biological remains from a Roman timber drain at 21 St Peters
Street, Colchester (site code: 2007.124)
Gary Andrew King and Allan Hall
Department of Archaeology, University of York, The King’s Manor, YO1 7EP
Summary
A subsample from a Roman timber drain was selected for detailed evaluation for its bioarchaeological
potential, primarily insect remains.
Both plant and insect taxa were present and in excellent
condition, though rather sparse. Analysis of the plant remains revealed the presence of both wild and
domestic occupation taxa including the presence of an exotic, fig. The insect fauna was largely
synanthropic in nature and resembled the indicator group associated with stable manure. Given the
context, the synanthropes are believed to be primarily background fauna suggesting the redeposition
of the material, most likely during the in-filling of the drain. The insect fauna also revealed some of
the earliest evidence for the presence of grain pests in Britain.
Keywords: COLCHESTER; ROMAN DRAIN; INSECT REMAINS; PLANT REMAINS
260
Evaluation of biological remains from a Roman timber drain at 21 St Peters
Street, Colchester (site code: 2007.124)
Introduction
In 2008, Colchester Archaeological Trust Ltd. excavated a nine metre long and 101.6 cm wide
timber drain at the 21 St Peter's Street site, within the town's Dutch Quarter. Using dendrochronology,
the construction of the drain has been dated to approximately AD 62, and the investigating field
archaeologists place the in-filling around 65-80 AD. Because of the presence on the site of some
deposits with waterlogged preservation (a very rare phenomenon in Roman Colchester), a 1.5 kg
subsample (Context 127) was submitted to the Centre for Human Palaeoecology, University of York
for evaluation of bioarchaeological potential, primarily through insect remains.
Methods
The sediment sample was inspected in the laboratory broadly following the procedures of
Kenward et al. (1980; 1985), for the recovery of plant and invertebrate macrofossils (three cycles of
admixture paraffin, 3 floatations). Plant and invertebrate remains in the resulting residue and washover
were recorded by ‘scanning’ using a low-power binocular microscope. Identification of insect remains
was carried out through comparison with material in the reference collection of the former
Environmental Archaeology Unit, University of York. Taxonomy and nomenclature for the insects
follow Kloet and Hincks (1977). Data were recorded on paper before being transferred to personal
computer.
Results
Context 127 (organic lowest fill of timber drain; silts sealed by in situ timber lid)
Sample 6 (1.5 kg sieved to 300 microns with paraffin floatation)
Moist, light-dark brown, stiff to crumbly, sandy-silt.
The washover yielded some mammalian bone fragments (a charred sheep ulna with coloration
suggesting firing temperatures around 700 degrees centigrade, an ungulate scapula, as well as ungulate
rib with evidence of butchery), eggshell, and oyster shell. Plant remains in the flot and residue both
consisted of ‘waterlogged’ seeds and fruits in a moderate state of preservation. The flot also contained
ample insect remains.
Most of the wild plant taxa recovered, including spike rush (Eleocharis sp.), lesser spearwort
(Ranunculus flammula L.), and Glyceria sp., are typical of wet places of various kinds. Orache
(Atriplex sp.) knotgrass Polygonum and docks (Rumex sp.) commonly inhabit disturbed ground. There
were a few taxa indicative of occupation and here, probably, domestic waste: traces of seeds of fig
261
(Ficus carica L.), fruitstone fragments of Prunus (sloe, plum, etc.) and nutshell fragments of Corylus
avellana L. (hazel). Some sclerotia (resting bodies) of the soil-dwelling fungus Cenococcum may
simply have arrived in imported soil or have formed from fungal mycelia that lived in the deposit at
some stage after formation.
The flot contained a relatively small number of insect remains. The fauna were primarily
synathropic (defined here as species associated with human occupation). The flot yielded one heavily
fragmented chrysomelid (leaf beetle) elytron, potentially representing a non-synanthropic species,
although this cannot be conclusively deduced due to the condition of the fossil. Additionally, the
presence of Phyllodrepa ?floralis/salicis could represent a nearby woodland environment or equally
be evidence of a more human-associated habitat through haystack refuse or stable dung (Koch 1989).
Given the context, it is also interesting to note the lack of aquatic invertebrates.
A high percentage (84 %) of the recovered insect remains consisted of synanthropic taxa,
presumably representative of the fauna of nearby buildings. Ptinus ?fur and Tipnus unicolor are both
characteristic of this category. While it has been found to inhabit bird nests, Ptinus fur is common in
mouldy straw and hay in barns and stables as well as cereal debris (Koch 1989). Tipnus unicolor is
found to frequent similar environments (Koch 1989) but is typical of older buildings. The recovery of
individuals of Lathridius minutus group and Gyrohypnus ?fracticornis is further evidence to support
the presence of mouldy decaying vegetation, particularly straw or hay (Böcher 1988; Koch 1989).
Although not necessarily indicative of the presence of hay or straw, Cercyon analis has been found in
decomposing plant debris and has been recovered from compost heaps and leaf litter (Hansen 1987).
Although Aphodius granarius has been recorded in rotting vegetation, the dung beetle is common in
stable manure heaps and may indicate the presence of foul matter.
While the drain fauna consisted primarily of facultative synanthropes (those forms most
commonly found in artificial environments but capable of surviving in nature), 27 % of the
synanthropic assemblage itself was contributed by strong synanthropes. The single individual of
Sitophilus granarius is evidence for the presence of cereal grains. S. granarius is capable of feeding on
damaged as well as undamaged grain, although it has been noted to have difficulty breaching husked
kernels. Cryptolestes ferrugineus is regarded as a secondary pest of cereals and is often found in grains
that have been worked or damaged. Palorus ratzeburgi is a scavenger of very spoiled grain and is
known to prey upon other grain pests. Both C. ferrugineus and P. ratzeburgi are also found in other
stored products, including flour, bran meal, and non-cereals such as dried fruit (Salmond 1957; Hunter
et al. 1973; Freeman 1980).
Discussion
Pests of stored products
262
One of the most interesting features of the Roman timber drain at 21 St Peter’s Street is the
presence of species associated with cereals and other stored products. Sitophilus granarius, the
granary weevil, is a common pest in granaries where both larvae and adults feed on whole cereals
(Hoffman 1986). S. granarius is considered a major pest of cereals and is noted to be very destructive,
resulting in considerable loss of stored grain. In the United Nations Food and Agriculture
Organisation’s report of 1947, it was suggested that 10 % of the world’s cereal production was lost to
insect attack; five decades ago 5 % of the loss was attributed to infestation by the granary weevil
(Munro 1966).
Whilst the granary weevil has been known to feed on grains in the early stages of spoilage
(Coombs and Woodroffe 1963), the other species present are often considered pests of cereals that
have been broken and become wet and mouldy, often as a result of attack by S. granarius. Observing
the natural succession of the infestation of stored grains, Coombs and Freeman (1955) have considered
species such as Cryptolestes ferrugineus and Palorus ratzeburgi to be secondary pests of stored
product cereals.
Although these stored product pests are believed to be able to overwinter successfully in the
unheated grain stores of Britain today as a result of the warmer-than-ambient temperatures existing in
the internal microhabitats (Solomon and Adamson 1955), the archaeological record indicates that they
were absent from Britain prior to the Roman invasion. Buckland (1978) proposes that this pre-Roman
absence is due to a combination of minimal importation of grain from the continent during the Iron
Age and the storage of grains in pits which would create a sealed carbon dioxide-rich environment
inhibiting infestation. The mass importation of cereals by the Roman army and civil administration as
well as the use of ventilated above-ground granaries may have enabled the pests to survive and
flourish.
The pre-Boudiccan deposits at One Poultry, London (Smith 2000) suggest that the species
entered Britain almost immediately after the Roman invasion. Moreover, having seemingly entered
Britain with the Romans, biogeographical mapping (c.f. King in press) suggests that the species spread
across England along with the Roman legions, entering the Roman Fort at the Millennium site at
Carlisle Castle by AD 72/3 (Smith and Tetlow n.d.) and the fort at Ribchester, Lancashire, by AD 71-4
(Large et al. 1994; Buxton and Howard-Davis 2000). Furthermore, with the Roman departure from
Britain, the granary beetles become notably absent from the record until the Norman Conquest.
At a minimum, the presence of the grain pests at the site in question here suggests the mass
storage of grains in the area and puts forth the possibility that the cereals may have been imported
rather than native.
Origin and deposition of material
263
Although the recovery of grain pests indicates the storage of grains near the site, they are not
necessarily evidence of the timber drain having serviced a granary, as was similarly proposed for the
Roman sewer in York (Buckland 1976). Kenward and Hall (1997) have also proposed that the
presence of grain pests along with ‘hay’ fauna, house fauna, and decomposers is characteristic of
stable manure, most likely equine. The grains would have served directly as a part of the mammals’
diet or, less possibly, the grain pests could have invaded residue grain in straw or chaff that was used
for bedding (Kenward forthcoming). Osborne (1983) demonstrated that insect fragments could
successfully pass through a human dietary tract without damage; it seems plausible that the same
would hold true for large non-ruminant herbivores.
An indicator group of organisms for stable manure is now recognised (Kenward and Hall
1997). From the invertebrates, stable manure can often be recognised through a combination of grain
pests, ‘hay’ insects, house fauna from the stables, and decomposers often associated with foul matter.
Along with the grain pests, the sample from 21 St Peters Street contained two commonly associated
house fauna taxa (Tipnus unicolor and Ptinus ?fur) and the dung beetle Aphodius granarius which is
strongly associated with stable manure. It also produced a range of fauna associated with plant debris,
particularly decaying hay and straw. This combination of fauna strongly supports the origin deposit as
stable manure.
While the presence of a stable manure indicator fauna in the timber drain could be indicative of
contemporaneous runoff and redeposition from the stable, the lack of aquatic insects supports the
possibility for in-fill or deliberate dumping as appears to be the case for the Roman deep wells at
Skeldergate and Bedern in York (Hall et al. 1980; Kenward et al. 1986).
Most of the plant remains were taxa likely to have been part of a local weed flora or to have
been imported with cut wetland vegetation (as litter for stables?), though with evidence from hazel nut
and fig for some material from domestic occupation. In the case of the fig, an exotic origin for the fruit
seems highly likely. The lack of evidence for cereals in a deposit containing grain pests is not
especially problematic since the routes by which such remains can travel on their way to a forming
deposit are complex (Hall and Kenward 1998).
Acknowledgements
I am grateful to Ben Holloway and Philip Crummy of the Colchester Archaeological Trust for
the opportunity to work on this material, to Harry Kenward for his assistance with this project and
comments on the draft, and to Terry O’Connor for assistance with identification of mammalian
fragments.
264
Appendix 2B:
Complete list of invertebrate remains recorded from the ‘detail’ recorded
subsample from the Roman timber drain at 21 St Peter’s Street, Colchester.
Order and nomenclature follow Kloet and Hincks (1964-77) for insects. Ecological codes used in
calculating statistics and minimum number of individuals (MNI) are given (they are explained in
Appendix 1C). The remains were of adults unless stated. ‘Sp.' indicates that record was probably an
additional taxon, ‘sp. indet.' that the material may have been of a taxon listed above it.
Taxon
Arachnida
Acarina sp.
Insecta
Diptera
Diptera sp. (pupa)
Coleoptera
Cercyon analis (Paykull)
Phyllodrepa
?floralis/salicis
Gyrohypnus ?fracticornis
(Muller)
Aleochara sp.
Aphodius granarius
(Linn.)
Tipnus unicolor (Piller &
Mitterpacher)
Ptinus ?fur (Linn.)
Cryptolestes ferrugineus
(Steph.)
Lathridius minutus group
(Linn.)
Palorus ratzeburgi (Wiss.)
Chrysomelidae sp. indet.
Sitophilus granarius
(Linn.)
Coleoptera sp.
Coleoptera (larvae)
Hemiptera
Psylloidea sp. (nymph)
MNI
Ecological Code
1
--
3
--
2
1
Rt
Rt
1
Rt
1
2
U
ob-rf
1
rd
2
2
Rd
G
1
Rd
4
1
1
G
-G
1
1
---
1
--
265
Appendix 2C:
Abbreviations for ecological codes used for interpretation of insect remains in text
and tables.
Lower case codes in parentheses are those assigned to taxa and used to calculate the group values
(the codes in capitals). Indivs - individuals (based on MNI); No - number.
No ‘certain’ outdoor taxa (oa) SOA
No strongly synanthropic taxa SSS
No ‘certain’ outdoor indivs NOA
No SS indivs NSS
No OA and probable outdoor taxa (oa + ob)
No uncoded taxa (u) SU
SOB
No indivs of grain pests (g) NG
No OB indivs NOB
No aquatic taxa (w) SW
No aquatic indivs NW
No damp ground/waterside taxa (d) SD
No damp D indivs ND
No strongly plant-associated taxa (p) SP
No strongly P indivs NP
No heathland/moorland taxa (m) SM
No M indivs NM
No wood-associated taxa (l) SL
No L indivs NL
No decomposer taxa (rt + rd + rf) SRT
No RT indivs NRT
No ‘dry’ decomposer taxa (rd) SRD
No RD indivs NRD
No ‘foul’ decomposer taxa (rf) SRF
No RF indivs NRF
No synanthropic taxa (sf + st + ss) SSA
No synanthropic indivs NSA
No facultatively synanthropic taxa
SSF
No SF indivs NSF
No typical synanthropic taxa SST
No ST indivs NST
266
Appendix 2D:
Complete list of plant remains and some other components of the residue from the subsample of
St Peters Street, Colchester.
All material was preserved by anoxic ‘waterlogging’ unless otherwise indicated. Nomenclature and
taxonomic order follow Tutin et al. (1964-80) for vascular plants. Abundance is presented using a
four-point semi-quantitative scale from 1—one or a few fragments or individuals (or a very small
component of the original sample volume) to 4—abundant remains or a large component of the
sample volume.
Name
Eleocharis sp.
Ranunculus flammula L.
Glyceria sp.
Atriplex sp.
Polgonum
Rumex sp.
Ficus carica L.
Prunus
Corylus avellana
Vernacular
Spike rush
Lesser spearwort
Sweet grass
Orache
Knotgrass
Docks
Fig
Sloe, plum, etc
Hazel
Abundance
2
2
1
2
1
2
1
1
1
267
Appendix 3
Stable-Isotopic Assays Relating to Modern Sitophilus granarius, Cereals, and
Quality Control Procedures
268
Appendix 3A:
Nitrogen-15 and Carbon-13 Results: Sitophilus granarius L.
Sample
Elemental
N
Mean δ15
NAIR
Elemental
C
Result δ13
CV-PDB
Mean δ13
CV-PDB
(%)
2.19
1.88
3.66
2.95
2.90
2.82
4.12
4.86
2.89
3.13
5.10
4.75
4.89
Result
δ15
NAIR
(‰)
1.71
1.70
1.08
0.57
-1.23
2.15
1.40
1.62
2.55
2.76
1.65
2.01
2.04
Ident.
S20091
"
S20092
"
S20093
"
S20094
"
S20095
"
S20096
"
S20097
"
S20098
(‰)
(%)
20.16
18.97
28.86
23.76
26.22
21.79
30.44
35.06
20.73
22.12
34.77
32.25
33.95
(‰)
-27.09
-27.21
-28.00
-27.98
-26.03
-24.04
-24.23
-24.56
-26.16
-26.39
-20.44
-20.96
-20.25
(‰)
"
4.01
3.07
2.56
29.46
-20.16
1.70
1.08
-1.23
2.15
1.51
2.65
1.83
269
-27.15
-27.99
-26.03
-24.04
-24.39
-26.27
-20.70
-20.21
Appendix 3B:
Nitrogen-15 and Carbon-13 Results: Cereals
Sample
Elemental
N
Result
δ15
NAIR
Mean δ15
NAIR
Elemental
C
Result δ13
CV-PDB
Ident.
(%)
(‰)
(‰)
(%)
(‰)
S20099
"
S200910
"
S200911
"
S200912
"
S200913
"
S200914
"
S200915
"
S00916
"
1.11
1.30
1.56
1.85
1.13
1.13
1.33
1.01
1.59
1.62
1.61
1.23
2.19
2.04
2.20
2.05
2.69
3.18
2.75
2.94
2.12
2.14
2.47
2.38
2.38
2.51
2.32
2.62
3.08
2.99
3.56
3.31
40.56
41.79
43.24
42.87
38.86
38.45
38.95
37.32
41.07
41.29
41.61
40.15
39.86
39.75
39.73
38.82
-27.59
-27.79
-28.30
-28.55
-24.94
-24.79
-24.50
-24.61
-26.21
-26.11
-26.49
-26.43
-22.63
-22.52
-21.95
-21.83
2.94
2.85
2.13
2.42
2.45
2.47
3.03
3.43
270
Mean δCV-PDB
13
(‰)
-27.69
-28.42
-24.87
-24.55
-26.16
-26.46
-22.57
-21.89
Appendix 3C:
Quality Control: Reference Standards Nitrogen-15 and Carbon-13
IA-R042
IA-R045
IA-R046
Bovine Liver
Ammonium
Sulphate
Ammonium
Sulphate
15
Mean
1 s.d.
n
Accepted
13
15
15
IAR005
Beet
Sugar
13
IA-R006
Cane
Sugar
13
δ NAir
(‰)
δ CV-PDB
(‰)
δ NAir (‰)
δ NAir (‰)
δ CVPDB (‰)
δ CV-PDB
(‰)
7.76
7.47
7.77
8.01
7.47
7.65
7.25
7.71
7.88
7.66
0.23
9
7.65
-21.53
-21.68
-21.61
-21.68
-21.69
-21.61
-21.58
-21.50
-21.63
-21.61
0.07
9
-21.60
-4.74
-4.64
-4.70
-5.15
-5.01
21.89
21.91
22.28
22.26
22.14
21.85
-26.08
-26.07
-26.19
-26.28
-26.20
-26.00
-26.03
-26.02
-11.83
-11.80
-11.86
-11.72
-11.79
-11.75
-4.85
0.22
5
-4.71
22.05
0.20
6
22.04
-26.11
0.10
8
-26.03
-11.79
0.05
6
-11.64
271
Appendix 3D:
Deuterium Results: Sitophilus granarius L.
Sample
Name
Sample
Descript.
Weight
Hydrogen
δ2HVSMOW
Mean
δ2HVSMOW
G1
G2
G3
G4
G5
G6
G7
G8
G9
G11
G12
G13
G14
G15
G16
G17
G18
G19
G20
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
SMOW
(BWB
Adjusted)
(‰)
-119.13
(mg)
0.67
(%)
2.26
(‰)
-116.21
0.52
3.07
-115.61
-118.51
0.47
2.70
-101.92
-104.48
0.77
2.62
-101.72
-104.27
0.84
2.47
-111.43
-114.22
0.81
2.13
-73.22
-75.06
0.90
2.91
-99.84
-102.34
0.85
2.47
-91.66
-93.96
0.80
3.55
-91.07
-93.36
0.57
2.46
-95.75
-98.15
0.30
4.50
-111.98
-114.79
0.67
2.49
-88.71
-90.93
1.30
2.05
-91.53
-93.82
0.47
2.53
-84.66
-86.78
0.98
2.08
-89.55
-91.80
1.35
2.17
-96.09
-98.50
0.78
2.15
-108.54
-111.27
---
---
---
---
1.63
1.65
-112.63
-115.45
272
(‰)
δ2HV-
Mean δ2H
(BWB
Adjusted)
(‰)
G21
Beetle
remains
0.83
2.22
-89.96
-92.22
G22
Beetle
remains
Beetle
remains
"
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
"
---
---
---
---
1.05
1.68
-96.23
-98.65
0.86
1.16
1.74
2.53
-93.77
-108.30
1.15
2.07
-101.41
-103.95
0.73
2.16
-98.36
-100.82
1.05
2.57
-111.24
-114.03
0.64
2.67
-97.67
-100.11
0.76
2.40
-90.52
-92.79
1.02
2.34
-99.34
-101.83
1.08
1.33
-107.16
-113.22
1.02
1.52
-101.24
1.08
1.43
-92.93
0.93
1.26
1.58
1.36
-94.04
-101.82
1.24
1.94
-83.26
-87.96
1.1
2.43
-97.99
-103.53
1.04
2.09
-105.46
-111.43
1.31
1.93
-93.93
-99.24
0.98
2.45
-89.67
-94.74
0.94
2.22
-94.93
-100.30
1.13
2.22
-93.40
-98.68
0.91
2.36
-89.19
G23
"
G24
G25
G26
G27
G28
G29
G30
G31
"
G32
"
G33
G34
G35
G36
G37
G39
G40
G41
"
Beetle
remains
"
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
Beetle
remains
"
273
-95.00
104.20
-96.12
-111.01
-106.96
-97.38
-110.09
-98.18
-93.49
-91.29
-99.36
-107.57
-94.23
-98.77
-96.45
Appendix 3E:
Deuterium Results: Cereals
Sample
Name
G42
"
Sample
Descript
.
Cereal
kernels
"
Weight
Hydrogen
SMOW
(%)
5.27
1.25
5.86
Cereal
kernels
0.93
5.43
"
"
0.90
5.25
G44
Cereal
kernels
Cereal
kernels
Cereal
kernels
"
1.10
5.06
1.06
5.09
1.20
5.02
1.10
5.03
Cereal
kernels
Cereal
kernels
1.10
5.58
0.97
5.18
"
"
1.17
5.03
G49
Cereal
kernels
0.92
5.59
"
"
1.13
4.94
G50
Cereal
kernels
Cereal
kernels
Cereal
kernels
1.08
5.15
1.06
5.42
1.06
5.12
G46
"
G47
G48
G51
G52
Mean
δ2HVSMOW
(mg)
1.18
G43
G45
δ2HV-
(‰)
87.68
99.33
102.4
9
101.0
5
70.86
78.87
87.21
88.02
97.98
124.0
5
127.7
9
113.0
6
118.5
1
81.91
86.44
59.65
(‰)
-93.51
δ2HV-SMOW
(BWB
Adjusted)
(‰)
-92.64
Mean δ2H
(BWB
Adjusted)
(‰)
-104.95
-98.80
-108.28
-101.77
-106.76
-107.52
-74.87
-83.33
-92.14
-87.62
-93.00
-92.57
-103.51
-131.06
-125.92
-135.02
-133.04
-119.45
-115.78
-125.21
-86.54
-91.33
-63.02
274
-122.33
Appendix 3F:
Deuterium Results: Water
Water
Sample Name
Water
δ2HVSMOW Mean δ2HVSMOW
(‰)
(‰)
Used to rinse samples
-47.43
-47.73
-47.58
Sample Description
275
Appendix 3G:
Quality Control Reference Standards: Deuterium
Check Sample
Replicate
IA-R002
(mineral oil)
1
2
3
4
5
6
7
8
9
10
IAEA-CH-7
(polyethylene)
BWB-II
(whale baleen)
δ2HV-SMOW δ2HV-SMOW (BWB Adjusted)
(‰)
(‰)
-112.81
-109.06
-107.51
-110.68
-110.59
-112.15
-111.60
-111.29
-110.78
-111.50
Mean
St. Dev.
-110.80
1.53
Accepted value
-111.2
1
2
3
4
5
6
7
8
9
10
11
12
-100.23
-101.11
-102.82
-102.32
-102.20
-101.64
-101.67
-101.89
-103.10
-100.70
-100.31
-101.78
Mean
St. Dev.
-101.65
0.92
Accepted value
-100.3
1
2
3
4
-106.64
-104.08
-103.05
-102.22
-109.31
-106.69
-108.88
-107.12
Mean
St. Dev.
-104.00
1.92
-108.00
1.29
Accepted value
-108
276
Appendix 3H:
Quality Control Reference Standards: Deuterium Water
Check Sample
Replicate
IA-R053
(natural water)
1
2
δ2HVSMOW
(‰)
-61.05
-62.39
Mean
St. Dev.
-61.72
0.95
Accepted value
-61.97
277
Appendix 3I:
Stable-Carbon and Nitrogen Bradford Analyses
Sample
Sit1
Sit2
Sit3
Sit4
Sit5
Sit6
Weight
1.03
1.54
1.23
1.55
1.18
0.97
%N
6.3
6.5
5.3
5.7
4.9
6.5
δ15N
5.61
5.70
7.00
6.28
6.52
6.27
%C
33.9
36.4
28.6
35.0
26.8
43.9
278
δ13C
-25.89
-26.05
-27.92
-27.89
-25.68
-25.75
C/N
6.26
6.56
6.31
7.12
6.40
7.93
Appendix 4
Stable-Isotopic Assays for Insect Remains Recovered from Erkelenz-Kückhoven,
Eythra, Plaußig, and West Stow
279
Appendix 4A:
Nitrogen-15 and Carbon-13 Results
Sample Elemental N Result δ-15NAIR Elemental C Result δ-13CV-PDB
Ident.
(%)
(‰)
(%)
(‰)
Sit27
1.22
5.85
10.44
-23.39
Sit28
0.45
1.06
4.77
-25.87
Sit29
1.67
4.66
13.61
-24.74
Sit30
3.22
-26.88
Sit31
9.37
-27.54
Sit32
0.96
2.73
8.39
-21.89
Sit33
4.88
-18.28
Sit34
Sit35
6.09
-24.62
Sit36
0.41
6.89
4.12
-22.96
Sit46
2.09
4.02
19.74
-25.16
Sit47
2.55
21.10
15.64
-24.76
Sit48
1.62
6.86
15.91
-27.29
Sit49
0.53
3.91
6.24
-27.39
Sit50
3.82
-1.86
33.91
-27.88
Sit51
1.13
-1.25
11.88
-26.09
Sit52
0.33
-6.02
4.22
-25.23
Sit53
0.35
-2.16
4.44
-26.25
Sit54
0.35
-5.99
4.14
-28.20
Sit55
4.73
-1.60
43.09
-25.42
Sit56
0.51
2.28
5.92
-29.38
Sit57
-
-
-
-
Sit58
9.30
10.10
46.35
-25.91
Sit59
1.65
3.09
11.56
-29.37
Sit60
-
-
6.66
-26.21
Sit61
2.49
0.80
15.64
-23.30
Sit62
0.48
-1.66
4.50
-25.65
280
Site
Plaußig
Plaußig
Plaußig
Plaußig
Plaußig
Plaußig
Eythra
Eythra
Eythra
Eythra
West Stow
West Stow
West Stow
West Stow
West Stow
West Stow
ErkelenzKückhoven
ErkelenzKückhoven
ErkelenzKückhoven
ErkelenzKückhoven
ErkelenzKückhoven
ErkelenzKückhoven
ErkelenzKückhoven
ErkelenzKückhoven
ErkelenzKückhoven
ErkelenzKückhoven
ErkelenzKückhoven
Appendix 4B:
Deuterium Results
Ident.
uncorrected
Result δ2
HV-SMOW
(‰)
BWB II
corrected
Result δ2
HV-SMOW
(‰)
Sit28
Sit32
Sit36
Sit46
Sit48
Sit49
Sit51
Sit52
-119.68
-106.18
-100.73
-84.46
-97.33
-84.47
-73.50
-123.49
-119.98
-106.45
-100.99
-84.68
-97.58
-84.69
-76.17
-123.81
correction
factor
1.0026
1.0026
1.0026
1.0026
1.0026
1.0026
1.0363
1.0026
Sit53
-121.46
-121.78
1.0026
-107.72
Sit54
-154.69
-160.31
1.0363
-104.21
Sit56
-125.88
-130.46
1.0363
-104.21
Sit58
-68.23
-70.71
1.0363
-104.21
Sit62
-101.07
-104.74
1.0363
-104.21
Sample
equilibration
281
mean
BWB II value
(‰)
Site
-107.72
-107.72
-107.72
-107.72
-107.72
-107.72
-104.21
-107.72
Plaußig
Plaußig
Eythra
West Stow
West Stow
West Stow
West Stow
ErkelenzKückhoven
ErkelenzKückhoven
ErkelenzKückhoven
ErkelenzKückhoven
ErkelenzKückhoven
ErkelenzKückhoven
Appendix 4C:
Quality Control Reference Standards: Deuterium
Mean
1 s.d.
n
Expected
IAEA-CH-7
BWB II
IA-R002
(PEF)
(Whale Baleen) (Miner