Environment, Insects and the Archaeology Of Egypt*
Eva Panagiotakopulu
University of Edinburgh
Paul Buckland
University of Bristol
Introduction
gypt lies in the overlap zone between two biogeographic realms, the Ethiopian (AfroTropical), which also includes the southern part of the Arabian Peninsula, and the Palaearctic, represented by the warm temperate zone of the Mediterranean. Apparent in its
now much depleted vertebrate fauna (cf. Manlius 2000; Manlius and Gautier 1999) and in
its flora, where Mediterranean elements extend down into Middle Egypt (Zahran and
Willis 1992), the overlap is also evident in its less well-studied insect faunas. Intuitively the
Nile provides a natural pathway for African elements to reach the Mediterranean and this
would have been more so during the early Holocene when stronger monsoonal circulation
made much of the present desert savannah (e.g. Haynes et al. 1989). Yet there are surprisingly few southern species in Egypt’s invertebrate fauna, which is overwhelmingly circumMediterranean in its affinities. Egypt has approximately 2,700 recorded species of beetle
(Coleoptera). Using the ground beetles, Carabidae, as an example, of the ~200 species recorded from Egypt, >95% are also found in Europe and the Near East, and a similar calculation for the water beetle families Dytiscidae and Haliplidae shows ~42 species of which
64% also occur in Europe. The scale of this overlap declines as distance from the Mediterranean coast increases, but Upper Egypt still shows affinities with the Palaearctic as well as
the Ethiopian Realm. Two major factors influence this pattern. One is the complex history
of the Nile and its floodplain, with periods during the Quaternary when its flow appears to
have failed completely (Said 1993; Butzer 1998). The other is the scale of destruction and
redistribution of biota by human activity.
No part of the Egyptian landscape can be viewed as ‘natural’ at the present day. The
scale of manipulation has accelerated in parallel with the country’s increasing population,
and it is doubtful whether anything approaching the ‘natural’ has existed since at the latest
the Roman period. Whilst increasing mid-Holocene aridity may have created the desert, its
present form is essentially capri-/ovigenic, and none of the neatly partitioned and irrigated
floodplain has escaped weeding, fertilising or more recently the application of insecticides.
Such impacts, from grazing pressure to storage loss and disease can potentially be traced
in changes in the insect fauna, and the frequent survival of fossil assemblages in archaeological contexts as a result of desiccation provides one source of this information. What are
E
*
This research began with the invitation to EP to take part in the Amarna project from Barry Kemp
to whom primary thanks are given. The work was partly funded by a major award from the
Leverhulme Trust to Jon Sadler in the Department of Geography at the University of Birmingham,
and additional funds were provided by the Egypt Exploration Society. The authors would like to
thank all members of the Amarna Project over the years and the several museum curators, who
have allowed EP to pry into their mummies, offerings and canopic jars. This paper has benefited
much from discussion with Peter Skidmore.
S. IKRAM & A. DODSON (eds), Beyond the Horizon: Studies in Egyptian Art,
Archaeology and History in Honour of Barry J. Kemp (Cairo, 2009).
PANAGIOTAKOPULU & BUCKLAND
lacking are long continuous sequences from the floodplain and Delta, without which we
have little idea of the nature of the riverine landscape at the beginning of settled agriculture. It is unclear whether wet, impenetrable African gallery forest extended as a finger
down to the Mediterranean forest zone preceding early settlement. Perhaps natural grazing
pressure, largely by elephant and hippopotamus, was sufficient to provide those clearings
into which early farmers could move, as aridity, and perhaps pastoralists, drove them off
the savannah. This has some bearing on Barry Kemp’s model (1989, fig. 7) of the early development of urbanization in the Nile valley, where his phase I can be preceded by a landscape of either forested or partly open floodplain. The debate over the nature of the preclearance landscape is an aspect currently under review in Europe consequent upon Frans
Vera’s recent re-evaluation (2002; cf also. Hodder et al. 2005) of the palaeoecological record, but mid-Holocene Egypt needs a similar discussion, once there is sufficient evidence.
To the east across Sinai, the Holocene pattern of forest and steppe is apparent from
work in Jordan and Israel (cf. Hunt et al. 2004; Robinson et al. 2006). Southwards Lake
Edward in Uganda (Beuning et al. 1997) has yielded a history of flow in the White Nile.
However throughout Egypt, despite the potential of the continually aggrading floodplain
and delta of the river, before the construction of the Aswan High Dam, no long cores have
been recovered and studied for their palaeoecological record.
The record of fossil insects from archaeological sites in Egypt has recently been reviewed (Panagiotakopulu 2001a), and Levinson and Levinson (1994) have also considered
the insect pests of stored products. Whilst this paper covers some of the same ground for a
different audience, it is also intended to point out directions in which future research
might move.
Archaeoentomology in Egypt
he involvement of scientific method in archaeology might be said to have had its origins in Egyptology, in the early nineteenth century vogue for the public display of
unwrapping of mummies (e.g. Pettigrew 1834). Both John Atkinson (1825) and Frederick
Hope (1834) reported the red-legged ham beetle, Necrobia rufipes (Deg.) (= N. mumiarum
Hope), from mummies, and Hope added two species of bacon beetle, Dermestes maculatus
Deg. (= D. roei Hope and D. elongatus Hope) and D. frischii Kug. (= D. pollinctus Hope) from
the head of a mummy from Thebes, brought to England by Wilkinson. His description of
the specimens as new species reflects the current state of taxonomy in the early nineteenth
century as zoologists got to grips with the naming of species. Hope (1842) took the study a
stage further in his examination of insects from the gut of a mummified sacred ibis, Threskiornis aethiopicus, a species last seen in Egypt in 1891 (Houlihan 1996), but present by the
tens of thousand in the animal necropolis at Tuna el-Gebel, where his specimen probably
came from (Kemp, pers. comm.). Armchair ornithologists had decided that the shape of
the beak precluded the bird feeding on insects, yet Hope’s dissection found the sacred
scarab, Scarabaeus sacer L., and two large tenebrionids, Akis reflexa and Pimelia pilosa F. in
the gut; T. aethiopicus was conclusively shown to be at least partly insectivorous. A later examination of a sacred ibis’ gut contents, by Blair (1935) added a carabid, Calosoma
chloristictum Klug, to the list of prey.
Ectoparasites were early noted on human mummies. The founder of palaeopathology,
Marc Armand Ruffer had detected nits and lice (Pediculus humanus Deg.) in the hair of
mummies in the Cairo Museum (Ruffer 1914). More recently, Joann Fletcher has noted
their high frequency in Old Kindom wigs made from human hair from Abydos (Fletcher
1994; 2000); Romans could be equally lousy (Palma 1991). Bed bugs (Cimex lectularius L.)
and human fleas (Pulex irritans L.: fig. 1) occur at New Kingdom Amarna (Panagiotakopulu
T
348
INSECTS, ENVIRONMENTS & ARCHAEOLOGY
and Buckland 1999; Panagiotakopulu 2001a). Lice are likely to have achieved a worldwide
distribution with their human host, and must share a long record of co-evolution (Reed et
al. 2007); they are also recorded from New World mummies (e.g. Horne 1979), but both
bed bug and flea probably had other primary hosts, the former probably in the roosts of
bats or pigeons (Panagiotakopulu and Buckland 1999), and the latter an early introduction
via pathways of gift exchange of furs from South America, where the primary host appears
to have been the guinea pig (Cavia porcellus Erxl.; Buckland and Sadler 1989; Dittmar and
Guillen 2003; Panagiotakopulu 2001a). The importance of insects as vectors in disease and
their impact on past societies has been the subject of much research, and popular as well
as academic publication (e.g. Zinsser 1935; Busvine 1976; Wolf et al. 2007). Panagiotakopulu (2004b) has recently reconsidered the origins and dispersal of bubonic plague in
terms of the biogeography of rats and fleas and the forced coincidence of introduced Indian black rat (Rattus rattus L.), endemic Nile rat (Arvicanthis niloticus L.) and people, which
resulted from the annual floods of the Nile. The hypothesis of a disease, present but not
fatal to the native rodent, being transferred to a new host and thereafter reaching plague
proportions in humans requires testing against the fossil record, both insect and small
mammal, and this is one of several examples where close cooperation between the archaeologist, archaeoentomologist and palaeopathologist will yield interesting results. Panagiotakopulu (2004b) has also commented upon the large numbers of Diptera in archaeological
deposits in Egypt and their role in the spread of Trachoma, river blindness, the frequency
of which in Minya in Middle Egypt the intrepid English traveller Amelia Edwards was
moved to remark upon (Edwards 1877).
It is perhaps not surprising that insects associated with stored meat products are
equally at home in mummies (Panagiotakopulu 2001; Strong 1981). At least one of these
species, D. frischii, occurs in the Neolithic of the Jura in the south of France (Ponel 1997),
although whether it is an indigenous circum-Mediterranean species, or had spread from
the East with the first farmers remains to be seen. Further north, in England, only two
outdoor species, D. lardarius L. and D. lanarius Ill., are known to be present by the Late
Bronze Age (Osborne 1989; Robinson 2000); the latter is now only a very rare import to
Britain. At least one widespread dipterous pest, the house fly (Musca domestica L.), which in
the north requires the artificial heat provided by man-made accumulations of animal dung
and other warm decaying litter, may be of Egyptian or Near Eastern origin (Skidmore,
pers. comm.; Panagiotakopulu 2004b). Common in samples from Egypt, by the Neolithic,
house flies were present at Thayngen-Weier at nearly 500m a.s.l. in Switzerland (Guyan
1981; Nielsen 1989) and in southern Sweden (Skidmore, pers. comm.).
The propensity of such pest species, both dipterous and coleopterous, to infest museum
specimens long after excavation has recently been highlighted by Buckland and Panagiotakopulu’s (2001) re-examination of the fauna and flora of the mummy of Rameses II, and
this might explain Attia and Kamel’s (1965) record of another bacon beetle (Dermestes carnivorus F.), from ‘pharaonic mummies’. Hinton (1945), following Fauvel (1889), regarded
this as a North American species, although Smith (1994) notes it as introduced to British
Columbia.
The problems of ascribing primary, natural distributions to now cosmopolitan species is
best approached from the fossil record. Despite their binomial names suggesting New
World origins, both the saw-toothed grain beetle (Oryzaephilus surinamensis L.), and the
lesser grain borer (Rhyzopertha dominica F.), have Palaearctic fossil records, which include
Egypt (Zacher 1934; Panagiotakopulu 1999). The former is also recorded from a charred
grain deposit from the Neolithic of Greek Macedonia (Valamoti and Buckland 1995), although it only becomes widespread in Roman deposits (Buckland 1991), whilst the latter,
whose thermal requirements suggest a warmer (Munro 1966), perhaps semi-arid tropical
349
PANAGIOTAKOPULU & BUCKLAND
origin, is present on Bronze Age Santorini (Panagiotakopulu and Buckland 1991), but may
not to have reached northern Europe until the late medieval period, where it is associated
with the rice weevil (Sitophilus oryzae L.) in an early fifteenth century pit in Southampton
(Grove 1995). The origins of the grain weevil (S. granarius L.), have been the subject of
much discussion (summarized in Buckland 1991). It is probable that its natural habitat lies
in the underground stores of wild grasses, including cereals, collected by rodents in the
Fertile Crescent, and that its transfer to human granaries took place as soon as storage of
plant foods became widespread. It is present in Old Kingdom Egypt (Solomon 1965; Howe
1972) at Saqqara, and had spread across Central Europe with the Linear Bandkeramik
(LBK), where it is known from two sites in Germany (Büchner and Wolf 1997; Schmidt
1998). Although relatively cold-hardy (Hunter et al. 1973), it only thrives outside the Mediterranean zone under conditions of large scale storage and transport, becoming almost
ubiquitous in the Roman Empire, occurring from a fort on the Antonine northern frontier
at Bearsden, 1 near Glasgow, to close to its desert frontier in Egypt at Mons Claudianus
(Panagiotakopulu and van der Veen 1997). Its presence in LBK deposits must be some indication of the nature of this expansion of early agriculture. Another probable primary
denizen of rodent burrows is the colydiid Aglenus brunneus (Gyll.) (cf. Dajoz 1977), which
Girling (1984) found in an otherwise natural Neolithic assemblage from the Sweet Track in
Somerset, England. At the present day, the species is strongly synanthropic, occurring in
mouldy plant residues in barns, stables and cellars, habitats more widespread during the
past (Kenward 1975; 1976); in the Roman period, this blind flightless beetle was accidentally taken with other grain pests to the remote quarry site of Mons Claudianus in the
Eastern Desert (Panagiotakopulu and van der Veen 1997).
Whilst the historical biogeography of the pests of stored products has its own intrinsic
interest, and may occasionally be used to support some curious hypotheses (cf. the discussion of Lasioderma serricorne (F.), the tobacco beetle, in Buckland and Panagiotakopulu
2001), their ubiquity in archaeological deposits in Egypt has other implications.
In any society where large scale transportation and storage of cereals forms an integral
part of organization, insects extract a tithe. Hoffman (1954) notes that one pest alone, S.
granarius (fig. 2), annually destroyed 5% of French cereal production before 1939, and
losses in less developed countries remain much higher. Haile (2006) quotes a number of
figures for various crops and pests, including village level storage losses for sub-Saharan
Africa of between 25 and 40%, and losses on individual sites may be virtually total, leading
to starvation. Problems of storage loss during the First World War led to a report recommending air tight storage, where the build up of carbon dioxide as the grain respires suffocates grain pests (Dendy and Elkington 1920). Buckland (1978) has argued that sealed pit
storage during the Iron Age in Britain precluded expansion of insect grain pests until the
establishment of centralized granaries after the Roman invasion. In the Mediterranean
other techniques either partly efficacious or palliative were necessary. It has been argued
that fire ash around saddle querns at Amarna functioned as an insecticide (Miller 1987),
and Panagiotakopulu and others (1995) have discussed the evidence for the use of a range
of natural plant and mineral insecticides, based upon evidence from Classical sources and
the well preserved charred assemblages of crops which had been stored in pithoi at Bronze
Age Akrotiri on Santorini. Whilst the presence of alien plant materials might be detected
in the routine study of plant macrofossil assemblages, if not interpreted, the use of substances such as tephra and diatomite is less easily observed. Both work by the cutting of
the thin tegumen between sclerites of the pest species leading to desiccation and death,
but the archaeological detection of such substances remains enigmatic.
1
Crowson’s identification, Sitophilus sp. in Dickson et al. (1989) is most likely to be this species.
350
INSECTS, ENVIRONMENTS & ARCHAEOLOGY
Archaeoentomological landscapes: potential and pit falls, the case of Pharaonic Amarna
n several ways, Amarna should be the ideal site for archaeoentomological research.
Desiccation has lead to near perfect preservation of insect chitin and the short period of
occupation, perhaps less than two decades in the middle of the 14th century BC on a pristine site on the desert edge between the Nile floodplain and the escarpment east of the
river, means that the problem of recycling of earlier materials in mudbrick structures and
deposits is minimal. Unfortunately the site has been extensively turned over and what escaped the early excavators has often been the quarry of treasure hunters. Well stratified
deposits, however, do occasionally survive, although perhaps potentially the best, the
bases of the numerous wells, are too deep – and therefore too expensive – to get at. In 1921
Eric Peet was excavating in the central part of the city on behalf of the Egypt Exploration
Society (Peet and Woolley 1923), and he examined a house that clearly had two phases.
The later had been constructed on the demolished remains of a previous house, itself
partly built into a shallow gravel pit in the desert surface. Remains of the doorposts of the
second house bore hieroglyphs that indicated that it had belonged to Ranefer, who had
charge of the Pharaoh’s horses. His house may have been built on a slight platform as a
mark of status, but more important from the palaeoecological viewpoint is the fact that the
older structure had been partly infilled with material other than building debris, and this
had been dug through by Peet’s workmen in the centre of the outer hall; elsewhere some
floors remained intact sealing the deposits. In 2003, Peet’s sections were cleaned back and
the house re-excavated for detailed planning. In the sections of the pit cut through the
house floor and accumulation beneath, down into the underlying sandy gravel of the
desert surface, it was evident that there were several irregular trampled surfaces within
layers of midden, gravel and other debris. The outer wall of the later house had been
constructed in a trench cut down through the deposits to a hard surface within the
accumulation, roughly at the level of the surrounding desert, and no obvious midden lay
above this level outside the structure. That man-made sediments extended deeper in
Peet’s pit indicated that both phases had been built into a pre-existing feature, probably a
shallow pit for the extraction of building materials, dug before the city had extended as far
to the south-east. The hard surface presumably related to the construction of the first
phase house on the site, but it is difficult to be certain whether the overlying material
relates to casual dumping of midden material into an abandoned property, something
which can be seen on deserted lots and derelict houses today in Minya, or to the deliberate
import of debris from a range of sources to make the platform on which the later house
stood. The frequency with which deposition was interrupted by clean gravel spreads, often
with a hardened surface, suggests that accumulation was intermittent and there can be no
doubt that at least the northernmost outer wall of the earlier house had been reduced to
its foundation course before midden accumulated over it. As well as a virtually complete
locust (fig. 3), the preserved insect evidence shows a relatively foul assemblage, dominated
by large numbers of the puparia of the housefly (Musca domestica L.), which had been
breeding in the rotting organic material, which appears to have included herbivore dung.
The insects do not resolve the question as to whether the debris was still foul, or had dried
out before deposition over the ruins of the early house. Despite its vernacular name, the
lesser mealworm (Alphitobius diaperinus Panz.), present in the deposits, would have
predated on the maggots of house flies, but the dispersed sclerites of individuals, rather
than the complete animals, including larvae, from the Workmen’s Village at Amarna
(Panagiotakopulu 1999a), suggest that the living population probably lay elsewhere,
although much invertebrate and vertebrate reworking is likely to have taken place in the
midden deposits. Cereal debris, including much straw and both charred and unburnt
I
351
PANAGIOTAKOPULU & BUCKLAND
grain, was evident throughout the deposit and examples of the grain weevil (Sitophilus
granarius L.) indicate that this was storage rather than field accumulation, although again,
as Osborne’s (1983) practical experiments in archaeoentomology have shown, these, along
with the more fragmented plant detritus, could have passed through either human or
animal guts. Scattered examples of the small flightless spider beetle (Gibbium psylloides
Czen.) may have strayed from its usual habitat, in foul rotting material, often faecal, but
very large numbers in a pot in the rubble filling beneath the southern part of the house
perhaps is indicative of partly dried excreta. There are taxonomic problems with the genus
Gibbium and older identifications, including the large numbers recorded from a New
Kingdom pot in the Berlin Museum by Zacher (1934), need to be checked for the congener
G. aequinoctiale Boield., but both species are characteristic of singularly foul residues, the
latter being recorded tunnelling into human faeces down a coal mine in England
(Constantine 1995).
Excavations at Amarna and adjacent Kom el-Nana have provided rare opportunities to
consider various other aspects of New Kingdom and Byzantine environments. This has
range from fish (Luff and Bailey 2000) to plant remains (e.g. Smith 2003), and has resulted
in several more synthetic studies (e.g. Kemp et al. 1994; Samuel 2000). Insect work had began with pig coprolites from the Workmen’s Village (Panagiotakopulu 1999a), but later
moved on to more general aspects of the environment of this part of the site (Panagiotakopulu 1999b). As well as the expected desert fauna, dominated by large tenebrionids,
what is apparent is the relative frequency of fodder and dung elements in the assemblages.
Their ubiquity goes beyond casual dispersal from the irrigated areas by the Nile, some 3
km away. All foodstuffs and water for both man and animal would have had to have been
brought up to the settlement, and casual inclusion of living and dead insects would have
been inevitable, but there remains the possibility that it was possible to grow some fodder
crops on the wadi floor at the time the site was occupied in the mid 14th century BC. The
insect evidence implies the cultivation of a legume, although a more varied mix is possible.
The frequency of desert tenebrionids in the relatively loose sand filling of archaeological features raises several taphonomic problems, not the least of which is when they actually ended up in the fossil record. Occasionally there can be no doubt, one specimen of
the large fossorial species Trachyderma cf. hispida from a charred deposit in the City was
complete, partly charred and intimately associated with the sediment. Other isolated
specimens are less secure and the need for recovery by the archaeoentomologist, aware of
the taphonomic implications, rather than casual collection by archaeologist during excavation is apparent. Where large numbers of the same species are found in an otherwise
empty pot – an example in the collections of the Fitzwilliam Museum in Cambridge has
over 100 specimens of Zophosis cf. carinata and there are similar assemblages from the
Workmen’s Village at Amarna – interpretation is more difficult, and it is probable that the
empty pot has functioned as a buried pit fall trap for the tunnelling insects over a considerable period of time. Levinson and Levinson (1996), influenced by the position of the
scarab (Scarabeus sacer L.) in Egyptian cult (cf. Alfieri 1956), interpreted large numbers of
the large, impressively spinose tenebrionid Prionotheca coronata Ol. in pots in Predynastic
tombs at Hu (Diospolis Parva) in Upper Egypt and from Early Dynastic Tarkhan in Lower
Egypt as evidence for ritual deposition, but it is much more likely that they represent accidental victims of empty vessels. Although the difference in species – none of these assemblages are mixed – must indicate differences in the environment, sufficient habitat data
have yet to be evaluated to elucidate this.
352
INSECTS, ENVIRONMENTS & ARCHAEOLOGY
A Byzantine Window
he most diverse insect faunas from an archaeological site in Egypt come from the Byzantine monastic site of Kom el-Nana, which lies towards the southern limit of Amarna,
north east of the modern village of Hagg Qandil, and now surrounded by its irrigated
fields. An aerial photograph, taken in the 1920s however, shows the mounds of the site
surrounded by desert, and it is only with the provision of modern irrigation methods that
agriculture has become possible in the immediate environs. Had the site been intermittently inundated, insect chitin would not have survived, and the fauna, which accumulated
in an abandoned room in the monastic complex, indicates cultivation in the immediate vicinity (fig. 4). Many of the fragments were packed together into accumulations suggestive
of faecal pellets, although it is difficult to suggest what animal, perhaps lizards. Much of
the assemblage is synanthropic, including the grain weevil, flour beetles (Tribolium spp.)
and the khapra beetle (Trogoderma cf. granarium Everts), the latter providing the earliest
fossil record of an important warm temperate to tropical pest of stored cereals, thought to
have originated in India or SE Asia (Peacock 1993).
What is particularly remarkable in the Kom el-Nana assemblages is the diversity of
dung beetles. In themselves, these suggest a different landscape from the present Egyptian
one, in that many require fresh dung deposited onto grazed pasture to complete their life
cycle. Extensive collecting around Amarna at the present day has produced only one dung
beetle, and suitable habitats have virtually disappeared from the intensively managed landscape. That extensive lush pastures, rather than intensive cultivation, by the Nile must
have existed in the New Kingdom is evident in tomb paintings. A fragment from the tomb
of Nebamun at Thebes (TTE2) in the British Museum, datable only a few years prior to the
Amarna Period, shows a large herd of cattle with their herdsman (Houlihan 1996: pl.xxii),
and similar themes appear in other wall paintings. Large, managed flocks of geese are also
shown in the same tomb. Whilst this is a long way from the Byzantine period, the pattern
is the same, one of extensive areas of productive pasture, rather than intensively cultivated,
individually ‘owned’ small fields, each farmer with his own cow, a buffalo or donkey for
traction, sustained wholly on fodder and crop waste, and all animal dung collected as fuel;
small numbers of sheep and goats are fed off the edge of the desert.
This encapsulates Egypt’s own ‘tragedy of the commons’ (cf. Harden 1968), enacted several times, perhaps from the apparent ease of the Arab Conquest to Mohammed Ali and
Nasser, if not before. Large estates have the luxury of extensive, less intensive agricultural
systems, which have space for elements in the natural biota, which compete with individual owners for resources, in this case dung, now extensively collected as fuel. It was these
estates that sustained the herds for temple sacrifice until the system fell apart in the
Graeco-Roman period, when grain production for a Mediterranean world took over (cf.
Bagnall 1993; Bowman 1990). In a ‘World System’, however, estates are not broken up for
the good of the fellaheen, but sequestered for cash crops, cotton or sugar cane monoculture in the nineteenth century, and the natural is driven out. Yet returning the land to the
‘people’ in small scale individual ownership is equally deleterious. Efforts to maximize
their output, cultivating up to the river, irrigation dike and desert edge, leads to destruction of any remaining natural environments and desertification, as the population pushes
towards the Malthusian breakpoint. Several times Egypt has reached this point in the past,
perhaps beginning with the collapse of the Old Kingdom, probably the result of disastrous
variation in the Nile flood. In the present landscape there is no space for wildlife, be it
dung beetle or gazelle, and the fossil record is our only window on the past. With sufficient sites, it will be possible to chart the progress of Egypt from riverine forest and savannah in the Early Holocene to the intensively managed irrigated agriculture and desert of
the Late Holocene.
T
353
PANAGIOTAKOPULU & BUCKLAND
BIBLIOGRAPHY
ALFIERI, A. 1956. ‘La véritable identité du scarabée sacré de l'Egypte pharaonique’,
Bulletin de la Société Entomologique d'Égypte 40: 451–52.
ATKINSON, J. 1825. [Letter concerning insects from Egyptian mummies], Transactions of
the Linnean Society, 14: 585–86.
ATTIA, R. and KAMEL, A. H. 1965. ’The fauna of stored products in U.A.R.’, Bulletin de la
société entomologique d'Egypte 49: 221–232.
BAGNALL, R. 1993. Egypt in Late Antiquity (Princeton: Princeton University Press).
BOWMAN, A. K. 1990. Egypt after the Pharaohs 332 BC–AD 642 (Oxford: Oxford University
Press).
BEUNING, K. R. M., TALBOT, M. R. and KELTS, K. 1997. ’A revised 30,000-year
paleoclimatic and paleohydrologic history of Lake Albert, East Africa’,
Palaeogeography, Palaeoclimatology, Palaeoecology 136: 259–79.
BLAIR, K. G. 1935. ‘Some ancient beetles from Egypt and Mesopotamia’, Proceedings of the
Royal Entomological Society of London 10: 19.
BÜCHNER, S. and WOLF, G. 1997. ’Der Kornkäfer – Sitophilus granarius (Linné) – aus
einer bandkeramischen Grube bei Göttingen’, Archäologisches Korrespondenzblatt
27: 211–20.
BUCKLAND, P. C. 1978. ‘Cereal production, storage and population: a caveat’, in S.
Limbrey and J. G. Evans (eds.), The effect of man on the landscape: the Lowland Zone
(London: Council for British Archaeology): 43–45.
— 1991. ‘Granaries, stores and insects. The archaeology of insect synanthropy’, in D.
Fournier, and F. Sigaut, (eds.), La preparation alimentaire des cereals (Rixenart:
PACT): 69—81.
BUCKLAND, P. C. and PANAGIOTAKOPULU, E. 2001. ‘Rameses II and the tobacco
beetle’, Antiquity 75: 549—56.
BUCKLAND, P. C. and SADLER, J. P. 1989. ’A biogeography of the human flea, Pulex
irritans L. (Siphonaptera : Pulicidae)’, Journal of Biogeography 16: 115—20.
BUSVINE, J. R. 1976. Insects, hygiene and history (London: Athlone Press).
BUTZER, K. W. 1998. Late Quaternary problems of the Egyptian Nile : stratigraphy,
environments, prehistory. Paléorient 23/2: 151—74.
CONSTANTINE, B. 1995. ‘Gibbium aequinoctale Boieldieu Ptinidae in coal mines: further
information’, Coleopterist 3: 87—90.
DAJOZ, R. 1977. ‘Coléoptères. Colydiidae et Anommatidae Paléarctiques’, Faune de
l'Europe et du Bassin Méditerranêen, 8 (Paris: Masson).
DENDY, A. and ELKINGTON, H. D. 1920. ‘Report on the effect of air-tight storage upon
grain insects’, Royal Society Grain Pest Committee Report 6.
DICKSON, J. H., DICKSON, C. A. and BREEZE, D. J. 1979. ‘Flour or bread in a Roman
military ditch at Bearsden, Scotland’, Antiquity 53: 47–51.
DITTMAR, K. and GUILLEN, S. 2003. ‘Studies on parts of the 28S rDNA on 1000 year
old fleas Pulex sp. from mummies of the Chiribaya Culture, southern Peru’ in N.
Lynnerup, C. Andreasen and J. Berglund (eds.), Mummies in a new millenium.
Proceedings of the 4th World Congress on mummy studies. Nuuk, Greenland, September
4th to 10th, 2001 (Copenhagen: Greenland National Museum and Archives and
Danish Polar Centre): 134–35
EDWARDS, A. B. 1877. A thousand miles up the Nile (London: Longmans, Green and Co.).
FAUVEL, A. 1889. ‘Liste des Coléoptères communes a l'Europe et a l'Amerique du Nord’.
Revue Entomologique de Caen, 8: 92–174.
FLETCHER, J. 1994. ‘A tale of hair, wigs and lice’, Egyptian Archaeology 5: 31–33.
— 2000. ‘Analytical techniques, Hair’, in Nicholson and Shaw (eds.) 2000: 499–500.
354
INSECTS, ENVIRONMENTS & ARCHAEOLOGY
GIRLING, M. A. 1984. ‘Investigations of a second insect assemblage from the Sweet
Track’, Somerset Levels Papers 10: 79–91.
GROVE, K. 1995. ‘A Study on Insect Remains in Archaeological Deposits’, in The
Archaeology of Chichester and District 1995 (Southern Archaeology and Chichester
District Council): 39–40.
GUYAN, W. U. 1981. ‘Puppen der Hausfliege, Puppen der “stallfliege”’, in W. U. Guyan,
Zur Viehhaltung im Steinzeitdorf Thayngen-Weier II: Archaologie der Schweiz, 4: 114–
16.
HAILE, A. 2006. ‘On farm storage studies on sorghum and chickpea in Eritrea’, African
Journal of Biotechnology 5: 1537–44.
HARDEN, G. 1968. ‘The tragedy of the commons’, Science 162: 124–42.
HINTON, H.E. 1945. A Monograph of the beetles associated with stored products, I (London:
British Museum [Natural History]).
HAYNES, C. V., EYLES, C. H., PAVLISH, L. A., RITCHIE, J. C. and RYBAK, M. 1989.
‘Holocene palaeoecology of the eastern Sahara: Selima Oasis’, Quaternary Science
Reviews 8: 109–36.
HODDER, K. H., BULLOCK, J. M., BUCKLAND, P. C. and KIRBY, K. J. 2005. Large
herbivores in the wildwood and modern naturalistic grazing systems (Peterborough:
English Nature).
HOFFMANN, A. 1954. ‘Coleoptérès Curculionides 2’: Faune de France (Paris: Lechevalier)
59: 487–1208..
HOPE, F. W. 1836. ‘Notice of several species of insects found in the heads of Egyptian
mummies’, Transactions of the Entomological Society of London 1: 11–13.
— 1842. ‘Observations on some mummied beetles taken from the inside of a mummied
ibis’, Transactions of the Royal Entomological Society of London 3: 191–93.
HORNE, P. 1979. ‘Head lice from an Aleutian mummy’, Paleopathology Newsletter 25: 7–8.
HOULIHAN, P. F. 1996. The Animal World of the Pharaohs (London, Thames and Hudson).
HOWE, R. W. 1965. ‘Sitophilus granarius L. Coleoptera, Curculionidae breeding in
acorns’, Journal of Stored Product Research 1: 99–100.
HUNT, C. O., ELRISHI, H. A., GILBERTSON, D. D., GRATTAN, J., MCLAREN, S.,
PYATT, F. B., RUSHWORTH, G. and BARKER, G. W. 2004. ’Early Holocene
environments in the Wadi Faynan, Jordan’, The Holocene 14: 921–30.
HUNTER, F. A., TULLOCH, J. B. M. and LAMBOURNE, M. G. 1973. ‘Insects and mites
of maltings in the East Midlands of England’, Journal of Stored Product Research 9:
119–41.
KEMP, B. 1989. Ancient Egypt. Anatomy of a civilisation (London, Routledge).
KEMP, B. J., SAMUEL, D. and LUFF, R. 1994. ’Food for an Egyptian City: Tell elAmarna’ in R. Luff, and P. Rowley-Conwy (eds.), Whither Environmental
Archaeology? (Oxford, Oxbow): 133–70.
KENWARD, H. K. 1975. ‘The biological and archaeological implications of the beetle
Aglenus brunneus Gyllenhal in ancient faunas’, Journal of Archaeological Science 2:
63–69.
—1976. ‘Further archaeological records of Aglenus brunneus Gyll. in Britain and Ireland,
including confirmation of its presence in the Roman period’, Journal of
Archaeological Science 3: 275–77.
LEVINSON, H. and LEVINSON, A. 1994. ‘Origin of grain storage and insect species
consuming desiccated food’, Anzeiger für Schädlingskunde, Pflanzenschutz,
Umweltschutz 67: 47–59.
355
PANAGIOTAKOPULU & BUCKLAND
— 1996. ‘Prionotheca coronata Olivier Pimeliinae, Tenebrionidae recognized as a new
species of venerated beetles in the funerary cult of pre-dynastic and archaic Egypt’,
Journal of Applied Entomology – Zeitschrift für angewandte Entomologie 120: 577–85.
LUFF, R. M. and BAILEY, G. N. 2000. ‘Analysis of size changes and incremental growth
structures in African catfish Synodontis schall schall from Tell el-Amarna, Middle
Egypt’, Journal of Archaeological Science 27: 821–36.
MANLIUS, N. 2000. ‘Historical ecology and biogeography of the hippopotamus in Egypt’,
Belgian Journal of Zoology 130: 59–66.
MANLIUS, N. and GAUTIER, A. 1999. ‘The wild boar in Egypt’, Comptes Rendus de
l'Academie des Sciences Serie III – Sciences de la Vie-Life Sciences 322: 573–77.
MILLER, R. 1987. ‘Appendix: Ash as an insecticide’, Amarna Reports IV: 14–16.
MUNRO, J. W. 1966. Pests of stored products (London: Hutchinson).
NICHOLSON, P. T. AND SHAW, I. (eds.) 2000. Ancient Egyptian Materials and Technology
(Cambridge: Cambridge University Press).
NIELSEN, B. O. 1989. ‘House fly puparia Musca domestica L. from a Neolithic field in
Thayngen-Weier, Switzerland’, Mitteilungen der Schweizerischen Entomologischen
Gesellschaft, 62: 5–8.
OSBORNE, P. J. 1983. ‘An Insect Fauna from a Modern Cesspit and its Comparison with
Probable Cesspit Assemblages from Archaeological Sites’, Journal of Archaeological
Science 10: 453–63.
— 1989. ‘Insects’, in P. Ashbee, M. Bell, and E. Proudfoot (eds.), Wilsford shaft excavations
1960–62 (London: English Heritage): 96–99.
PALMA, R. L. 1991. ‘Ancient head lice on a wooden comb from Antinoë, Egypt’, JEA 77:
194.
PANAGIOTAKOPULU, E. 1999a. ‘An examination of biological materials from coprolites
from XVIII Dynasty Amarna, Egypt’, Journal of Archaeological Science 26: 547–51.
— 1999b. Insect remains from Tell el Amarna: interim report. File Report, Amarna
Project, Cambridge.
— 2001a. ‘Fleas from pharaonic Amarna’, Antiquity 75: 499–500.
— 2001b. ‘New records for ancient pests: archaeoentomology in Egypt’, Journal of
Archaeological Science 28: 1235–46.
— 2004a. ‘Pharaonic Egypt and the origins of plague’, Journal of Biogeography 31: 269–275.
— 2004b. ‘Dipterous remains and archaeological interpretation’, Journal of Archaeological
Science 31: 1675–1684.
PANAGIOTAKOPULU, E. and BUCKLAND, P. C. 1991. ‘Insect pests of stored products
from Late Bronze Age Santorini, Greece’, Journal of Stored Product Research 27: 179–
84.
— 1999. ‘The bed bug, Cimex lectularius L. from Pharaonic Egypt’, Antiquity 73: 908–911.
PANAGIOTAKOPULU, E., BUCKLAND, P. C., DAY, P. M., SARPAKI, A. and
DOUMAS, C. 1995. ‘Natural insecticides and insect repellents in Antiquity’, Journal
of Archaeological Science 22: 705–10.
PANAGIOTAKOPULU, E. and VAN DER VEEN, M. 1997. ’Synanthropic insect faunas
from Mons Claudianus, a Roman quarry site in the Eastern Desert, Egypt’ in A.
C.Ashworth, P. C.Buckland and J. P.Sadler (eds.), Studies in Quaternary Entomology
– An Inordinate Fondness for Insects. Quaternary Proceedings 5: 199–206.
PEACOCK, E.R. 1993. Adults and larvae of hide, larder and carpet beetles and their relatives
Coleoptera: Dermestidae and of Derodontid Beetles Coleoptera: Derodontidae (London:
Royal Entomological Society of London).
PEET, T. E. and WOOLLEY, C. L. 1923. The City of Akhenaten I (London: EES).
356
INSECTS, ENVIRONMENTS & ARCHAEOLOGY
PETTIGREW, T. J. 1834. A History of Egyptian Mummies and an account of the worship and
embalming of the sacred animals by the Egyptians with remarks on the funeral ceremonies
of different nations and observations of the Canary Islands, of the ancient Peruvians,
Burman priest &c. (London: Longman, Rees, Orme, Brown, Green and Longman).
PONEL, P. 1997. ‘Sucession des assemblages de Coléoptères à Chalain 3’, Monographies de
Chalain et Clairvaux, 3: Que Chalain 3, le Néolithique final à la fin du Ive millénaire:
113–18.
REED, D. L., LIGHT, J. E., ALLEN, J. M. and KIRCHMAN, J. J. 2007. ‘Pair of lice lost or
parasites regained: the evolutionary history of anthropoid primate lice’, BMC
Biology 57: doi.10.1186/1741–7007–5–7.
ROBINSON, M. 2000. ‘Middle Mesolithic to Late Bronze Age insect assemblages and an
Early Neolithic assemblage of waterlogged macroscopic plant remains’, in S. P.
Needham (ed.), The passage of the Thames. Holocene environment and settlement at
Runnymede (London: British Museum Press): 146–67.
ROBINSON, S. A., BLACK, S., SELLWOOD, B. W. and VALDES, P. J. 2006. ‘A review
of palaeoclimates and palaeoenvironments in the Levant and Eastern
Mediterranean from 25,000 to 5000 years BP: setting the environmental background
for the evolution of human civilisation’, Quaternary Science Reviews 25: 1517–41.
RUFFER, M. A. 1914. ‘Pathological notes on the royal mummies of the Cairo Museum’,
Mitteilungen zur Geschichte der Medizin und der Naturewissenschaften und der Technik
13: 239–48.
SAID, R. 1993. The river Nile. Geology, hydrology and utilization (Oxford: Pergamon Press).
SAMUEL, D. 2000. ‘Brewing and baking’ in Nicholson and Shaw (eds.) 2000: 537–76.
SCHMIDT, E. 1998. ‘Der Kornkäfer Sitophilus granarius Schön. Curculionidae aus der
Schuttschicht des bankeramischen Brunnens von Erelenz-Kückhoven. Öst Rhein’,
Amt für Bodendenkmalpflege Hg. Brunnen der Jungteinzeit. Internat. Symposium
Erkelenz 27–20 Okt. 1997 Materialen zur Bodendenkmalpflege 11: 261–69.
SKIDMORE, P. 1992. ‘Notes on the taxonomy of the puparia of British Sphaeroceridae’,
Dipterists' Digest 13: 6–22.
SMITH, R. 1994. ‘Effects of Alien Insects and Microorganisms on the Biodiversity of
British Columbia's Insect Fauna’, in L. E. Harding and E. McCullum (eds.)
Biodiversity in British Columbia. Our changing environment (Canadian Wildlife
Service): 190–219.
SMITH, W. 2003. Archaeobotanical investigations of agriculture at Late Antique Kom el-Nana
Tell el-Amarna (London: EES).
SOLOMON, M. E. 1965. ‘Archaeological Records of Storage Pests: Sitophilus granarius L.
Coleoptera, Curculionidae from an Egyptian Pyramid Tomb’, Journal of Stored
Product Research 1: 105–107.
STRONG, L. 1981. ‘Dermestids – an embalmer's dilemma’, Antenna 5: 136–39.
VALAMOTI, S. M. and BUCKLAND, P. C. 1995. ’An early find of Oryzaephilus
surinamensis from final Neolithic Mandalo, Macedonia, Greece’, Journal of Stored
Product Research 31: 307–309.
VERA, F. W. M. 2002. ‘A park-like landscape rather than a closed forest’, Vakblad
Natuurbeheer, special issue Grazing and grazing animals 2000: 13–15.
WOLF, N. G., PANOSIAN DUNAVAN, C. and DIAMOND, J. 2007. ‘Origins of major
human infectious diseases’, Nature 447: 279–83.
ZACHER, F. 1934. ’Vorratsschädlinge und Speicherwirtschaft im alten und neuen
Ägypten’, Forschung und Fortschritte 10: 347–48.
ZAHRAN, M. A. and WILLIS, A. J. 1992. The Vegetation of Egypt (London: Chapman and
Hall).
357
PANAGIOTAKOPULU & BUCKLAND
ZINSSER, H. 1971. Rats, lice and history (London: Bantam Sociology).
358
INSECTS, ENVIRONMENTS & ARCHAEOLOGY
FIG. 1
Fleas (Pulex irritans L.) from the Workmen’s Village, Amarna.
FIG. 2
Charred wheat from deposits beneath the floor of Ranefer’s house, Amarna. The grain
shows evidence of intense infestation by the weevil Sitophilus granarius L.
359
PANAGIOTAKOPULU & BUCKLAND
FIG. 3
Complete specimen of the migratory locust (Schistocerca gregaria Forskål) from the midden
deposit beneath Ranefer’s house, Amarna.
FIG. 4
Insect remains from the Byzantine monastery at Kom el-Nana.
360