Six-legged Hitchhikers: An Archaeobiogeographical Account of the Early
Dispersal of Grain Beetles
Gary A. King1,*, Harry Kenward2, Edith Schmidt3, and David Smith4
Abstract - Grain-associated insect species are economically important and archaeologically significant. Their dispersal
around the globe and eventually across the North Atlantic region surely occurred through human transport rather than
naturally. Most beetle cereal pests are now nearly cosmopolitan in their distribution, but their prehistoric ranges appear to
have been more restricted. What is known or surmised of the early dispersal of these insect species is summarized, and the
role of archaeobiogeographical data in investigating past human contact evaluated. Analysis of fossil and historic records
of grain-associated beetles suggests that their dispersal corresponded with assumptions concerning human movement and
interaction in the past. There is a significant fossil record for some grain beetles, but it is incomplete and predominantly
from northwest Europe. More fossils are needed from across the Palaearctic and North Africa. The examination of preagricultural
natural deposits in the Middle East, North Africa, and the Indian Subcontinent might reveal the original ranges
of the pest species, the stages by which they entered into association with humans, and their earliest dispersal. With a more
complete fossil record, the grain fauna may provide a useful proxy by which to evaluate cultural contact and human migration
into the North Atlantic region in the past.
1Department of Archaeology, Durham University, South Road, Durham DH1 3LE, UK. 2Department of Archaeology, University
of York, The Kings Manor, York YO1 7EP, UK. 3Archaeoentomology, Eco Concept and Pictures Consultants, Gerda-
Weiler Str. 10, Freiburg 79100, DE.4Department of Classics, Ancient History and Archaeology, University of Birmingham,
Birmingham B15 2TT, UK. *Corresponding author - gking500@googlemail.com.
Introduction
Grain-associated insect pests play an important
role in the reduction of human food resources in
the present day (e.g., McFarlane 1989, Payne 2002,
Tyler and Boxall 1984). Of these species, beetles
(Order Coleoptera) are arguably the most socioeconomically
significant. Additionally, beetles have
been a critical factor leading to food depletion in
the past (e.g., Fitch 1879, Kirby and Spence 1859,
Munro 1966). These synanthropic beetles are of
archaeological, biogeographical, and ecological
importance as they are species that were dispersed
alongside humans (often well beyond their naturally
viable distributions), and thus can be used as secondary
evidence of past human movement and/or trade
(cf. King 2010b). But where did they originate, and
when did they spread? How significant were they to
past economies as people moved across the North
Atlantic?
Dispersal pathways by which organisms spread
between areas can be classified into corridor, filter,
and sweepstakes routes (Cox and Moore 2000). In
a corridor route, suitable habitats exist between the
source and invaded areas. The majority of organisms
would be able to disperse between the areas
with little difficulty. A filter pathway presents a more
limited range of habitats, so that only organisms that
can exist in those habitats can disperse between the
regions. The end regions in sweepstake dispersal are
“islands” surrounded by a “sea” (sometimes literally)
of unsuitable habitat. Depending on the scale at
which habitats are examined, the dispersal of storage
pest species can be placed in any one of these categories,
but they are usually seen as undergoing the
third type of dispersal, by hopping between isolated
islands of artificial habitat (mainly food stores in
temperate areas).
Elton (1958) introduced the concept of human
activity as a mechanism for the passive distribution
of animals and plants beyond their natural geographic
ranges. Buckland (1981, 1990) attempted to
trace the dispersal of stored-product pests by people
on the basis of the archaeoentomological records
available at the time. He successfully demonstrated
that pests were transported in the past, but patterns
of movement or origins for the evaluated species
could only be guessed at in view of the paucity
of fossil data, especially from beyond the British
Isles. Buckland offered speculations and urged for
more archaeoentomological investigations across
Eurasia. The themes introduced by Buckland have
subsequently been touched on and amplified by others
(e.g., King 2010b; Panagiotakopulu 2000; Plarre
2010; Smith and Kenward 2011, 2012). Three decades
after Buckland’s seminal 1981 paper, we use
the currently available fossil and literary evidence to
revisit Buckland’s review.
2014 Journal of the North Atlantic No. 23:1–18
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G.A. King, H. Kenward, E. Schmidt, and D. Smith
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The Archaeological Record of Storage Pests
Many thousands of samples from archaeological
deposits have now been examined for insect
remains, and although most have been from sites in
the British Isles, a substantial number have come
from Scandinavia, mainland Europe, the Middle
East, and North America. These insect remains are
usually preserved under anoxic conditions by “waterlogging”,
and while typically disarticulated into
individual sclerites, or groups of sclerites, they may
be in superb condition, often bearing scales or setae,
and sometimes containing male genitalia. Charred
fossils are moderately common and represent the
main source of information in more arid areas.
Storage pests appear early in the Holocene fossil
record in the Middle East and North Africa, and their
subsequent spread across Europe and into North
America can be traced, albeit with some large gaps
in time and space. For Britain, we have a fairly full
record for the past two millennia, from which patterns
of invasion, abundance, and species composition
can be followed.
Good as the archaeological record often is, caution
is essential in using it to trace the early history of insect
species. We need to recognize that there are problems
related to the presence of intrusive remains that
can occur, post-dating the deposit in which they are
found. Contamination of samples during excavation
and storage, and during processing in the laboratory,
have all been documented (Kenward 2009). There is
also a danger of misidentification of fossils, which
may be in poor condition, or of simple errors in recording
and data handling. All of these have certainly
occurred, so that special care is needed in evaluating
the significance of published rare records of insects
from unexpected periods. Unfortunately, some remains
that (with hindsight at least) were clearly contaminants
have been recorded from archaeological
associations without comment; some other records
are at least suspect. Were every archaeological record
to be accepted unquestioningly, various Australasian
beetles would now be regarded as originating in Europe,
for example. Ideally, any suspect remains should
be re-examined, but many have probably been lost.
Insect remains from tombs need to be approached
with particular caution, for recent invasion, especially
during transport and museum storage, is not just possible,
but highly likely, and not all such contaminants
will appear recent.
The Grain Beetles
The beetle fauna associated with stored cereals
includes, in addition to native species, a group
of species which have been ecologically classified
by Kenward (1997) in the context of northwest
European archaeology as “strong synanthropes”,
i.e. a species group comprising taxa that are mostly
thermophilous and generally dependent on artificial
habitats for survival in the region. However, as the
classification is climate-dependent, certain species
are able to survive beyond the boundaries of the human-
created artificial environments in other regions.
Information on the commonly recovered grain fauna
is presented in Table 1.
Based on their ability to infest or attack undamaged
cereal kernels, grain beetles can be classified
as primary or secondary pests. Primary pests are
capable of successfully attacking, feeding, and
multiplying on undamaged grains and often complete
their entire development within a single grain
(Semple et al. 1992). From archaeological contexts
in temperate regions, four primary beetle pests
of stored grains have been commonly recovered:
the granary weevil, Sitophilus granarius; the rice
weevil, S. oryzae; the maize weevil, S. zeamais;
and the lesser grain borer, Rhyzopertha dominica.
Most of these species have been recovered from
non-synanthropic habitats, primarily in tropical
and subtropical areas, and seem to be favored by
high temperatures for completion of their development
(see Table 1). In contrast, the flightless
species S. granarius has yet to be found outside of
human-created environments and is able to adjust
to unheated indoor conditions in Northern Europe;
indeed the granary weevil is not favored by high
temperatures (perhaps indicating an origin in an
upland area; see King, in press c; Plarre 2010).
Secondary pests are not capable of attacking
previously undamaged grains and do not complete
their development within a single grain. Beetle species
in this category tend to attack a wider range
of commodities than primary pests (Semple et al.
1992). Several of the commonly recovered secondary
pests of archaeological interest, e.g., beetles of
the genera Oryzaephilus, Cryptolestes, and Palorus,
are thermophilous and may have originally
exploited a habitat of loose bark and fungoid wood.
Buckland (1990) posited that moulds were the link
between the natural and artificial environments; the
damp grain can mimmick the microhabitat of fungoid
bark.
Some of the species that are generally listed
among “pests” in the stored-products literature,
e.g., the biscuit beetle, Stegobium paniceum, 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. Similarly, the large
scavenger beetles, e.g., Tenebrio spp. and Blaps
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G.A. King, H. Kenward, E. Schmidt, and D. Smith
Table 1. Common grain-associated beetle fauna from archaeological contexts. The optimum range of temperature for each species is given in parentheses following the range of temperatures it
is known to survive.
Species Common name Habitat Temperature range (optimum)
Alphitobius diaperinus (Panzer) Lesser mealworm Attacks dried animal and plant material (Rees 2007) 16–35 °C (30–33 °C)
Cryptolestes ferrugineus (Stephens) Flat or rust grain beetle Found in wheat, rape, maize, meal, and flour as well as dried fr uit (Horion 1960) 18–42.5 °C (33–35 °C)
Cryptolestes turcicus (Grouvelle) Mediterranean flat beetle Found in cereals and cereal products as well as dried fruit (Ha lstead 1993) 19–37 °C (30–35 °C)
Gnatocerus cornutus (Fabricius) Broad-horned flour beetle Attacks dried animal and plant material, especially grains and cereal products (Rees 2007) 16–32 °C (24–30 °C)
Oryzaephilus surinamensis (L.) Saw-toothed grain beetle Attacks a wide range of commodities including cereal and cereal products, dried fruit, and 20–38 °C (30–33 °C)
nuts (Koch 1989)
Palorus ratzeburgii (Wissman) Small-eyed flour beetle Attacks stored cereal products, particularly mouldy grain resid ues previously attacked 18–37 °C (30–32.5 °C)
by primary pests (Brendell 1975)
Rhyzopertha dominica (Fabricius) Lesser grain borer Attacks cereal grains, especially wheat, barley, rice, and sorghum (Rees 2007) 19–38 °C (32–34 °C)
Sitophilus granarius (L.) Granary weevil Attacks a range of stored products, especially cereal grains, a nd nuts (Hoffman 1954) 15–34 °C (26–30 °C)
Sitophilus oryzae (L.) Rice weevil Attacks a range of stored cereals including rice, rye, maize, w heat, barley, and millet (Harde 1984) 18–35 °C (26– 31 °C)
Sitophilus zeamais Motschulsky Maize weevil Attacks a range of stored cereals including rice, rye, maize, w heat, barley, and millet (Harde 1984) 18–35 °C (26– 31 °C)
Stegobium paniceum (L.) Biscuit or drugstore beetle Attacks a range of farinaceous foods (Buck 1958) 15–34 °C (25– 28 °C)
Tenebroides mauritanicus (L.) Cadelle Feeds principally on dried material of plant origin, especially grains and cereal products (Rees 2007) 15–38 °C(28–30 °C)
Tribolium castaneum (Herbst) Rust-red flour beetle Attacks stored grains and cereal products (Brendell 1975) 20–42 °C (32–37.5 °C)
Tribolium confusum Jacquelin du Val Confused flour beetle Infests stored grains and cereal derivatives (Brendell 1975) 19–37.5 °C (30–34 °C)
Trogderma granarium Everts Khapra beetle Attacks dried material animal and plant origin, including grain s and cereal products (Rees 2007) 24–43 °C (33–37 °C)
spp., are regularly found with stored products but
have also been noted in nature, such as in rotting
bark and birds’ nests, and feed on a range of organic
matter (see Koch 1989). Many other “pests” had a
wider range of habitats in the past and are best just
regarded as domestics in the context of archaeology
(“house fauna” as defined by Kenward and Hall
1995; see also Carrott and Kenward 2001, Kenward
and Carrott 2006). Notable among these domestics
are the various spider beetles of the genera Ptinus,
Tipnus, Gibbium, and Niptus. Other components
of the “house fauna” which are now often found in
storage habitats include various latridiids and cryptophagids,
Mycetaea subterranea (F.) (Endomychidae),
and Typhaea stercorea (L.) (Mycetophagidae),
all mould feeders.
Records of Storage Fauna in Space and Time
Setting the stage: The earliest records
The archaeological as well as documentary
evidence suggest that the principle grain pests first
formed an association with human beings in the
Middle East, but the origin of some species may
have been much further east (King 2010b, in press
c; Plarre 2010). The archaeoentomological record
for Eastern Asia is very limited. However, recent
work by Obata et al. (2011) has revealed the earliest
evidence for stored-product pests. Obata and associates
discovered impressions of Sitophilus zeamais
in early Jomon potsherds dating to ca. 9000 BP from
the Sanbonmatsu site in Kagoshima Prefecture,
Japan (Obata et al. 2011). Although recorded as S.
zeamais, the species-level identification was based
solely upon the length of the individuals and the
claim that adult maize weevils are on average larger
than adult rice weevils. However, size is not reliable
in separating the two species, and genetic analysis
is often necessary (see Hidayat et al. 1996); as such,
the identifications may be safer if conservatively
considered as S. zeamais/oryzae. 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.
As an aside, there are later records from the
Eastern Asia. “Maize” weevil fossils have also been
recovered from contexts at Fujiwara Palace (ca. 8th
century AD) and Kiyosu Castle (ca. 16th century AD)
(Mori 2001), and impressions have been noted from
Late Jomon pottery in Kyushu (Yamazaki 2005). Chu
and Wang (1975) reported Trogoderma persicum
(Pic.) (i.e., T. variabile Ball.) and Sitophilus oryzae
(i.e., oryzae or zeamais) from a tomb dated about
2100 BP in Hunan Province in China. The oldest writ2014
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G.A. King, H. Kenward, E. Schmidt, and D. Smith
4
ten account of grain pests in China, the Ĕryă, dates
between the 5th and 2nd centuries BC (Karlgren 1931).
In the Middle East, G.A. King (unpubl. data) has
noted holes that resemble the characteristic damage
made by Sitophilus weevils in charred wild einkorn
from the Natufian era Abu Hureyra I site in modern
Syria, ca. 13,000 BP. (However, bubbling during
charring can produce similar damage; Kenward et
al. 2008.) Interestingly, the damaged einkorn grains
were recovered from a sample containing acorns,
perhaps furthering the hypothesis that acorns were a
pathway for the introduction of the species into artificial
habitats (Buckland 1981, Howe 1965, Zohary
1969). The earliest fossils of grain pests in the Middle
East came from layer VI at Hacilar, SW Anatolia,
dated to 7700–7550 BP, in the Pre-Pottery Neolithic
C period (Helbaek 1970). Helbaek described charred
fragments of several adult Sitophilus in small heaps
of charred wheat and barley. Additionally, one of the
grains contained an adult weevil, and some kernels
showed evidence of lengthwise tunnelling. As early
evidence of S. oryzae and S. zeamais has yet to be
recovered in the region, the unidentified Sitophilus
are likely to have been S. granarius. More conclusive
evidence for the presence of the granary weevil
in the region at the time came from a near-contemporaneous
well at Atlit-Yam, Israel (ca. 7500 BP)
where Kislev et al. (2004) recorded 27 specimens of
Sitophilus granarius.
Moving substantially forward in time, the Harra==
Hubullu Tablets XI–XV provide the earliest
written zoological account of stored-product species,
listing 33 names of crop and stored-product
pests, e.g., uh.še.kú and uh.zí(d).da (Landsberger
1934; see also King 2010b). While believed to
have been compiled during the 9th century BC in
bilingual Sumero-Akkadian script, the tablets are
thought to be derived from Hammurabian period
lists (ca. 3792–3750 BP), which were in turn developed
from even older ones (Harpaz 1973). The
grain pests are also mentioned in other literature
of the period, for example, the Sumero-Akkadian
proverb “A piece of linen is spread for a flea, a tissue
for a moth, a granary for grain pests” (Bodenheimer
1947). Helbaek (1970) referred to weevilravaged
grain, providing indirect evidence for the
presence of S. granarius in Assyrian and Hellenistic
barley from Nimrud. Hopf and Zachariae (1971)
recorded the grain weevil in grain deposits dating
to around 3000 years ago from Tel Arad in the
Northern Negev, Israel. Kislev and Melamed (2000)
discovered insect remains of about the same date
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
in association with charred wheat, Triticum
parvicoccum. Other storage pests were also noted,
including: Alphitophagus bifasciatus (Say), Oryzaephilus
surinamensis (adult and pupa), and an adult
and whole larva of Tenebroides mauritanicus.
As with the Sumero-Akkadians, the ancient
Egyptians left a scant documentary record of grain
pests. Egyptian inscriptions include images of some
invertebrates that can be identified to genus (cf.
Harpaz 1973, Levinson and Levinson 1998), but the
ancient Egyptian language, like biblical Hebrew, apparently
lacked a comprehensive term for “insect”
(King 2010b). The Ebers Papyrus is an Egyptian
medical document (ca. 3552 BP) describing magical
formulae and remedies and is one of the earliest written
records containing methods for controlling pests.
The Ebers Papyrus XCVIII contains instructions
for deterring kkt-animals using burnt gazelle dung
diluted in water. The kkt-animals may be a reference
to grain weevils (Panagiotakopulu et al.1995).
However, kkt transliterates as small animal, and the
species identification is purely speculative, based on
context (King 2010b).
The earliest archaeological accounts for Egyptian
grain pests were made by Helbaek (cited in Solomon
1965), who reported Sitophilus granarius from the
ca. 4900 BP Tomb in Saqqarah. Solomon (1965) also
mentions that S. granarius was recovered from the
tomb beneath the Step Pyramid of Saqqarah, ca. 4300
BP. Sitophilus granarius and Stegobium paniceum
were recovered at the tomb of Queen Ichetis at Saqqarah,
ca. 4334–4150 BP (Chaddick and Leek 1972).
Several specimens each of Trogoderma granarium
and Stegobium paniceum (the earliest fossil
evidence for both), were identified in a wheat deposit
from a Middle Kingdom tomb at el-Gebelein
(4181–4055 BP) (Panagiotakopulu 2003). A Tribolium
species was noted in a mid-3rd millennium BC
(5000–4000 BP) Egyptian tomb by Alfieri (in Andres
1931). Tribolium confusum was reported from
an offering pot from ca. 3000 BP (Alfieri 1976).
Zacher (1937) recorded T. castaneum from Egypt
ca. 3500 BP. Rhyzopertha dominica and S. paniceum
were present in Liverpool Museum collections from
Twelfth Dynasty Kahun, 3990–3800 BP (Panagiotakopulu
1998). R. dominica was recovered from a
small sample of barley and is the earliest on record;
it was also noted in a botanical sample from a vessel
in Tutankhamun’s tomb, ca. 3345 BP (Alfieri 1931),
while Zacher (1937) recorded T. castaneum, S. paniceum,
Oryzaephilus surinamensis, and R. dominica
from another vessel from the tomb.
Samples from the Workmen’s Village at Tell el-
Amarna (thought to be dated between 3350–3323
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G.A. King, H. Kenward, E. Schmidt, and D. Smith
BP on pottery) yielded grain pests in probable
pigsty deposits—fossils of Sitophilus granarius and
Rhyzopertha dominica were recovered. Moreover,
Panagiotakopulu (1999) discusses the remains of S.
granarius and Palorus ratzeburgii from coprolites at
the site. Panagiotakopulu (2001) lists Tribolium confusum,
T. castaneum, Palorus subdepressus (Wollaston)
and Cryptolestes turcicus from Pharoanic
Amarna. Zacher (1934a, b) noted Oryzaephilus surinamensis
(L.) from a Minoan period vessel, 3350
BP. Panagiotakopulu and Buckland (2010) provide
a recent review of insect remains from Egyptian
archaeological sites.
Moving westward: records from prehistoric
Europe
How soon the grain pests were carried to Europe
is uncertain. The earliest evidence of cereal pests in
the region comes from bandkeramic wells in Eythra
village in the Leipzig region of Germany from Well
2 (cal. radiocarbon dated 7269–7180 BP) and Well 1
(dendro-date 7034 BP) (Schmidt 2005a). The oldest
records for the golden spider beetle, Niptus hololeucus
(Fald.), and for Gibbium psylloides (Czemp.)
are reported from the bandkeramik well in Eythra
(Leipzig) (Schmidt 2005a). Large numbers of granary
weevils have also been recorded from the well
at Plaussig, near Leipzig, dated dendrochonologically
to 7219 BP (Schmidt, in press a) and from a
well at Erkelenz-Kückhoven near Cologne dendrodated
7040 BP and 7007 ± 5 BP (Schmidt 1998, in
press b) (Fig. 1). Büchner and Wolf (1997) have also
recorded S. granarius from an underground pit, at
Göttingen, Germany, dated 6030 BP.
Populations of the granary weevil were well established
in central Europe less than 500 years after
the earliest known fossil appearance of the species
in the Middle East. However, the pathway that it
followed is unclear, although it was surely introduced
via human migration or exchange rather than
natural dispersal (see King 2010b for discussion).
The only other Neolithic evidence for S. granarius
comes from a cast in a piece of pottery from Servia
(6700 BP) in south Macedonia, Greece (Hubbard
1979).
Although the fossil record for the granary weevil
in Neolithic Europe is sparse, it is even more limited
for the other stored-cereal pests. A single charred
head of Oryzaephilus surinamensis was reported
from Mandalo in western Macedonia, Greece, dating
5490 ± 55 BP (Kotsakis et al. 1989, Valamoti
and Buckland 1995). Additionally, the cadelle Tenebroides
mauritanicus has been recovered from
Erkelenz-Kückhoven (Schmidt 1998, in press b) and
Plaussig (Schmidt, in press a) in Germany (Fig. 1).
Tenebroides mauritanicus has also been noted from
Figure 1. Illustrations of intact insects from Reitter (191 1) and Jamestown photos from Keng et al. (2010).
2014 Journal of the North Atlantic No. 23
G.A. King, H. Kenward, E. Schmidt, and D. Smith
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the Singen-Offwiese, Germany, which is associated
with the Groß Gartach culture dendro-date 6950 BP
(Dieckmann et al. 1997, Schmidt 2007).
The fossil evidence of grain pests continues to
be limited for the Bronze and Iron Ages. Sitophilus
granarius was found in a Middle Bronze Age site
in Northern Italy (Fasani 1976), and in Late Bronze
Age France at the site of Lake Bourget (Pecreaux
2008). Moreover, Middle Bronze Age contexts at
Cova Punta Farisa in Fraga Huesca, Spain yielded
the remains of Rhyzopertha dominica (Alonso
Martinez and Buxo Capdevilla 1993). The biscuit
beetle Stegobium paniceum has been recovered from
sites dating to Late Bronze Age Britain: Runnymede
Bridge, Staines, Surrey (Robinson 1991) and
Wilsford, Wiltshire (Osborne 1989). Chowne et al.
(1986) reported the biscuit beetle from an Iron Age
site in Britain (Tattershall Thorpe, Lincolnshire). S.
granarius, R. dominica, and Tribolium sp. have been
recovered from Siriguarch, Alcañiz, Teruel, Spain
(Compte and Perales 1984). There is a record of
Tenebrio molitor L. from prehistoric Britain, dated
4000 BP (Howard et al. 1999), notable in view of its
rarity in the later fossil record.
The northern Mediterranean shore
While the grain pests almost certainly spread
into Europe via the Mediterranean shores, early
fossil records from that area do not yet exist; the
contrast in known archaeological evidence of grain
pests from southern compared to central Europe is
probably at least in part a result of different cultural
and preservational circumstances (particularly the
lack of known waterlogged deposits in the south).
There is a modest amount of evidence from later
periods.
The fossil record for insects from pre-Roman
Greece is remarkably limited. A carbonized specimen
of Sitophilus granarius was recovered from a
sample of barley at the “unexplored” mansion complex
at Knossos, with a Late Minoan date, around
3425 BP (Jones 1984). Shaw and Shaw (1995)
noted S. granarius and Tribolium confusum from
a contemporaneous site at Kommos. Panagiotakopulu
and Buckland (1991) identified the remains
of S. granarius, Rhyzopertha dominica, Stegobium
paniceum, and Oryzaephilus sp. in samples from the
West House, Akrotiri Santorini, Thera, ca. 3500 BP.
In literature, the Greek terms κίς, κορίς, ϕθείρ, σής,
ϊψ, σκυίψ, and θρίψ were used to refer to small insect
pests, and Κίς, in particular, is believed to have been
associated principally with insects that infest grains
or pulses (Beavis 1988, King 2010b).
Expanding to the Atlantic seaboard: The Roman
Empire in Europe
There is abundant documentary evidence for
the Roman period, which dates back to Cato’s De
Re Rustica, ca. 235 BC (Ag. XCII). The Roman
authors express particular concern over a grain
pest called curculio—an insect that was capable of
ravaging enormous heaps of grain (e.g., G. Virgil
I.CLXXXVI). The term was even used by Plautus
to portray a greedy, gluttonous, and unscrupulous
character (PC. 219–221). Several authors also provide
insight into the construction of granaries and
methods for preventing contamination of cereals by
curculio (e.g., Arch. Vitruvius VI.VI.IV, RR. Varro
I.L.VII; RR., Columella I.VI.XV). Both Varro and
Columella depict curculio as a primary pest of stored
cereals, capable of infesting undamaged grains, and
King (2010b) suggests that the term may have been
applied to both Sitophilus granarius and Rhyzopertha
dominica, with the former being more commonly
associated.
Fossil records for the early Roman period are
rare. However, Cappadocia, Thrace, Carthaginian,
and Oscensian districts in Hither Spain, and Apulia
adopted pest control measures against infestations
of curculio (RR. Varro I.L.VII). In the absence of
an established fossil record for 36 BC, Varro’s text
affords the best evidence for the distribution of
curculio at that time, implying that the species was
established along most of the Mediterranean coast
of southern Europe during the Roman Republic (see
Carr 1838, SG. Strabo VI.III in King 2010b).
Later Roman sites have yielded abundant fossil
records of the grain pests, the frequency of recovery
apparently reflecting the intensity of research, so
that the archive for Britain greatly exceeds that for
mainland Europe.
Fossil records from the first century AD. The earliest
evidence of Roman-age grain pests (i.e., Oryzaephilus
surinamensis and Sitophilus granarius), dating
to 30 AD, was reported from Neuss (Novaesium)
in Germany (Cymorek and Koch 1969, Koch 1970).
Knörzer (1970) also identified the rice weevil S. oryzae
in a sample containing charred rice from early
first century AD contexts from Neus-Novaesium
IV, Germany. From Touffréville Calvados, France,
Ponel et al. (2000) reported S. granarius, Stegobium
paniceum, Tenebrio obscurus, and Oryzaephilus sp.,
dated ca. 75 AD. From beneath the AD 79 tephra
at Herculaneum, Naples, Italy, Dal Monte (1956)
noted larval, pupal, and adult S. granarius as well as
a single Oryzaephilus sp. in infested charred wheat.
Additionally, Sitophilus granarius was recovered
from first-century AD contexts at Alphen aan den
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G.A. King, H. Kenward, E. Schmidt, and D. Smith
Rijn, Netherlands (Kuijper and Turner 1992) and
Valkenburg fort (Hakbijl 1988).
The grain pests appear to have entered Britain
following the arrival of the Roman legions. It is
now three decades after Buckland’s (1981) review,
and there are still no records to support the presence
of grain pests in Britain prior to the arrival
of the Roman military forces (King 2010b, Smith
and Kenward 2011). At the Poultry site in central
London, a number of buildings, workshops, yards,
and pits dating to just after 47 AD—the start of the
Roman occupation—and sealed by the 60 AD “fire
horizon”, interpreted as the burning of London
during the Boudiccan revolt, have yielded myriad
insect remains, including: Sitophilus granarius,
Oryzaephilus surinamensis, Cryptolestes ferrugineus,
Palorus ratzeburgii, Alphitobius diaperinus,
Tenebrio molitor L., and Tenebroides mauritanicus
L. (Rowsome 2000, Smith 2011). Similarly,
deposits at 20–30 Gresham Street, London, dated
to 50–75 AD, produced large numbers of individuals
of C. ferrugineus, S. granarius, A. diaperinus,
O. surinamensis, and P. ratzeburgii (Smith and
Tetlow 2004). The same range of species was identified
from a timber drain at 21 Saint Peters Street,
Colchester, constructed immediately following the
Boudiccan revolt (King and Hall 2008).
The grain pests arrived in northern England in the
later part of the first century AD. The Roman Fort at
the Millennium site at Carlisle Castle, dated before
and shortly after 72/73 AD, yielded Cryptolestes
ferrugineus, Sitophilus granarius, Oryzaephilus
surinamensis, Tribolium castaneum, and Palorus
ratzeburgii (Smith 2010). Grain pests have been
noted at other sites in and near the Roman fortress at
Carlisle (Kenward and Carrott 2006, Kenward et al.
2000), and the fort at Ribchester, Lancashire 71–74
AD (Buxton and Howard-Davis 2000, Large et al.
1994). King (2010b) identified O. surinamensis,
C. ferrugineus, P. ratzeburgii, Tenebrio obscurus,
and S. granarius from a late first-century AD cesspit
at Spurriergate, York. Similarly, 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, including: Tenebroides mauritanicus,
C. ferrigineus, O. surinamensis, P. ratzeburgii,
T. obscurus, and S. granarius—representing a massive
infestation (Hall and Kenward 1976, Kenward
and Williams 1979).
Second, third, and fourth centuries AD. The
grain pests continue to be represented in France,
Germany, and England beyond the first century. A
Gallo-Roman granary in Amiens, France, which was
burned during the second century, yielded a fauna
which included Stegobium paniceum, C. ferrugineus,
O. surinamensis, Tenebrio sp., P. ratzeburgii, and
Sitophilus granarius (Matterne et al. 1998, Yvinec
1997). S. granarius also continued to be present in
the second century of Hambacher Forest (deposit
Hambach 512), NW Cologne in Germany (Schmidt
2006). Grain-associated insect species have been
recorded from a number of sites throughout England
(e.g., Buckland 1982; Kenward 2009; Kenward and
Carrott 2006; Robinson 2002, 2003; see also Smith
and Kenward 2011) and have been found as far north
as Invereskgate in West Lothian, Scotland (Smith
2001, 2004) until the end of the fourth century AD.
The second century provides more evidence of Tribolium
castaneum (Hall and Kenward 1990, Hall
et al. 1980), as well as the arrival of A. diaperinus
(Hall and Kenward 1990; Kenward and Allison
1995; Kenward et al. 1986, 2000) and Stegobium
paniceum (Hall and Kenward 1990) in Northern
England. Furthermore, P. subdepressus (Osborne
1971), C. turcicus, and Tribolium confusum (Girling
1983) appear to have been introduced to England
by the third and fourth centuries AD. South of the
Mediterranean, Oryzaephilus sp. and C. turcicus
have been recovered from a second-century quarry
site at Mons Claudianus in Egypt (Panagiotakopulu
and van der Veen 1997).
The later Roman period also provides the earliest
archaeological evidence for the movement of grain
pests as stowaways in ships. Pals and Hakbijl (1992)
recovered Sitophilus granarius, O. surinamensis,
C. ferrugineus, P. ratzeburgii, Tenebrio molitor,
Alphitophagus bifasciatus Say, 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. Sitophilus sp. (probably S. granarius)
was reported from a third-century ship wreck near
Guernsey (Rule and Monaghan 1993). The presence
of cereal pests in these vessels shows that grain was
still a traded commodity in the outer reaches of the
Roman Empire during the second and third centuries,
and that it was insect contaminated.
A curious rarity of grain pests: The early medieval
period
The early medieval period (5th–11th centuries
AD) is considered as a unit because there are no
clearly authentic British records of the main grain
pests from it (and incidently because Tipnus unicolor
(Piller and Mitterpacher), characteristic of many
Roman sites, was rare, or more often absent; see
Kenward 2009, Kenward and Whitehouse 2010).
2014 Journal of the North Atlantic No. 23
G.A. King, H. Kenward, E. Schmidt, and D. Smith
8
Both the grain pests and T. unicolor were present
in a few samples from Anglo-Scandinavian Coppergate,
York, but there are good reasons to regard
them as quite probably contaminant, including their
rarity, sometimes their preservational condition,
and the regrettable fact that samples from sites rich
in grain pests were processed concurrently in the
same laboratory (Kenward and Hall 1995:760–762).
Tipnus unicolor was, however, rather abundant at an
Early Christian rath site of Deer Park Farms, Antrim,
Northern Ireland (Kenward et al. 2011). While not
considered to be a typical stored-product pest, T.
unicolor is part of the “house fauna” species group
and is generally an indicator of long-lived, high-status
buildings, with proportions of this spider beetle
believed to increase with the improved cleanliness
of buildings (Kenward 2009). Kenward and Hall
(2000) reported the presence of eggs of the cestode
genus Hymenolepis in Anglo-Scandinavian deposits
from Walmgate, York. The life cycle of Hymenolepis
is known to be able to include grain pests, particularly
Tenebrio and Tribolium, as intermediate hosts
(see King and Henderson, in press).
No grain pests were found in numerous samples
from 11th-century Viborg, Denmark, though Ptinus
raptor (Sturm) and P. fur (L.) (the latter a common
species in stored products) were present (Kenward
2005a, b). Similarly, the distinctive P. raptor
has been identified from the Viking-Age site in
Kaupang, Norway (Barrett et al. 2004, 2007). From
the same Norwegian site, Buckland et al. (2001)
recorded Ptinus remains as P. pusillus (Sturm) and
P. palliatus (Perr); however, neither was found in
the subsequent study. These investigations underline
the rarity or absence of the classic pests in northwest
Europe during the early medieval period. However,
a Merovingian grave near Pattonville, Ludwigsburg,
SW Germany, from the late 6th to the middle of the
7th century exhibited large numbers of C. ferrugineus
found in a byzantine bowl (Bofinger and Ebinger-
Rist 2009, Schmidt 2010). From the coffin of the
Ottonian queen Editha (died 946), which was found
in Magdeburg, Germany, Schmidt (2012) recovered
large numbers of C. ferrugineus, Sitophilus granarius,
P. fur, and Tipnus unicolor. By the 9th century,
the large scavenger beetles appear to have arrived
in Ireland; Reilly (2003) reports Tenebrio molitor/
obscurus from Essex Street West site in Dublin. A
contemporaneous site, Wood Quay in Dublin, also
yielded Blaps lethifera Marsham (O’Connor 1979).
In contrast to the lack of convincing records from
northern England, Sitophilus granarius, O. surinamensis,
C. ferrugineus, and T. obscurus were present
in small numbers in late 10th- and early 11th-century
deposits at the Poultry site (Smith 2011). However,
deposits at this site appear to have been extensively
reworked in the past, and so these fragments could
be intrusive from later medieval deposits. The same
explanation could be advanced for the single individual
of S. granarius recovered from deposits of a
similar age at the Guildhall site, London (Smith and
Morris 2008). Alternatively, perhaps this find was a
result of differences in trade links, the demands of
London being great enough to necessitate at least
occasional imports of foreign grain, but further
investigation of tightly dated and securely sealed
waterlogged deposits of the period is desirable.
Emerging beyond the sphere of prior Roman
influence: The later medieval to early modern
periods
There are numerous records of grain pests from
Britain after the Norman Conquest, especially of
S. granarius, O. surinamensis, and C. ferrugineus,
set against a background of generally declining
urban faunal diversity. Amongst the pests, there
is—subjectively—a broad tendency for S. granarius
to become relatively more abundant, perhaps
because grain was cleaned better (it is hard to sieve
out weevils, both because of their larger size which
more closely approximates the size of the grains they
infest, and their presence inside the grains). Roman
sites, such as those at Ribchester, Carlisle, and York,
mentioned above, typically yield assemblages in
which S. granarius is appreciably less abundant than
C. ferrugineus, and especially O. surinamensis (for
example, at Tanner Row, O. surinamensis, C. ferrugineus,
and S. granarius were in the approximate
ratio 5: 2: 1; Hall and Kenward 1990). By contrast,
S. granarius may be proportionally much more
abundant in later deposits, such as the post-medieval
site at Coffee Yard in York, where there were more
than twice as many S. granarius as O. surinamensis,
and no records of other storage pests apart from
spider beetles, probably best seen as domestics at
this site (Robertson et al. 1989). At another site in
York, The Bedern, grain pests were present in pitfill
deposits dated 13th to early 17th century, but only
O. surinamensis and S. granarius were found, in
roughly equal numbers (Hall et al. 1993). It is worth
noting that the medieval deposits at The Bedern also
yielded the eggs of the tapeworm Hymenolepis sp.
(Hall et al. 1993).
A similar trend towards predominance of S. granarius
may have occurred in Germany, although the
evidence so far is slight: a large and varied fauna
from 15–16th century Neuss included a range of
domestic and storage beetles, but only S. granarius
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G.A. King, H. Kenward, E. Schmidt, and D. Smith
a medieval ditch at Iveagh Market (Reilly 2001) and
a 16th-century context from George’s Quay, Limerick
(O’Donovan 2002).
Crossing the Atlantic: Records from Iceland,
Greenland, and North America
Numerous insects, mostly synanthropes, were
carried across the Atlantic to Iceland and Greenland
(e.g., Forbes et al. 2010, Konráðsdóttir 2007, Sadler
1991, Sveinbjarnardóttir et al. 2007). Among them
were occasional storage pests, though none seem to
have become established, which is hardly surprising
in areas where cereals could rarely be grown successfully.
There are a few records from Iceland. Oryzaephilus
surinamensis was found at late medieval Holt
(Sveinbjarnardóttir 1983). Both Sitophilus granarius
and O. surinamensis were found in post-medieval
deposits in a high-status farm at Reykholt, Iceland,
indicating storage of grain, though the grain may
have been imported (Sveinbjarnardóttir et al. 2007).
Sitophilus granarius was present at post-medieval
Stóraborg (Buckland et al. 1992, Sveinbjarnardóttir
et al. 1981), and post-medieval layers at Bessastaðir
contained O. surinamensis and S. granarius
(Amorosi et al. 1992). Remarkably, both Sitophilus
and Oryzaephilus—surely carried in supplies of
grain—have been found in later Norse deposits, apparently
pre-dating the mid-14th century, at Nipaatsoq
in Greenland (Panagiotakopulu et al. 2007).
It is hardly surprising that a range of Old World
synanthropes arrived in North America with provisions
carried by the European colonists as well as
in the lively trade that followed. Evidence for this
importation is particularly notable in the recently
examined well fill deposits dated 1611–1617 from
James Fort, Colonial Jamestown, Virginia, USA,
which have revealed a range of invasive stored product
species, e.g., O. surinamensis, Sitophilus granarius,
S. oryzae/zeamais, C. ferrugineus, Palorus
sp., B. lethifera, Alphitophagus bifasciatus, Typhaea
stercorea (L.), and Mycetea subterraneus (Marsh.),
as well as native fauna, i.e., cf. Lasioderma serricorne
(F.) (King et al. 2010) (Fig. 1). Additionally,
a privy used at the English settlement at Ferryland,
Newfoundland, Canada, dated 1621–1673, yielded
a fauna with a range of European synanthropes,
including O. surinamensis and S. granarius (Bain
and King 2011, Bain and Prévost 2010, Prévost and
Bain 2007). Both S. granarius and Tenebrio obscurus
were recovered from a mid-17th-century shipyard
in Québec City in southeastern Canada (King, in
press b). Analysis of late 17th-century deposits in
Colonial Boston produced more assemblages rich
in imported insects, among them Tenebroides mauritanicus,
Oryzaephilus sp., Gnatocerus cornutus
among the classic grain pests (Koch 1970). This site
was unusual for its records of Ptinus latro (F.), P. cf.
clavipes (Panzer), and Niptus hololeucus (Fald.) as
well as the commonly encountered P. fur (L.) and
Tipnus unicolor. In a 14th-century cesspit in Constance
(Germany), S. granarius and O. surinamensis
were found in great numbers (Schmidt 2005b).
There are two records of the rice weevil S. oryzae
from northern England during this period: one from
early modern Hull (Carrott et al. 1995), and the
other from a 19th-century drain fill in the center of
York (Hall et al. 2006), in the latter case together
with R. dominica. The presence of insects in sieved
samples from the wreck of an 18th-century AD merchantman
off the Red Sea coast of Egypt at Sadana
Island is implied, and a jar with an insect-proof neck
described (Ward 2001).
Beyond the sphere of prior Roman influence, the
grain pests found their way to Scandinavia and Ireland.
A single S. granarius was recorded from a cesspit
fill dated 1275–1300 in Oslo, Norway (Kenward
1988), and the granary weevil was also present in a
medieval context from the Lofoten Islands (Vågon),
Norway (Buckland and Panagiotakopulu 1995).
From medieval Gothenburg, Sweden, Andersson
(1992) reported S. granarius and Tenebrio obscurus.
The stored-product pests, S. granarius and P. fur,
were recovered from the remains of a 13th-century
shipwreck of the frame-timbered vessel Oskarshamn
off the coast of Sweden (Lemdahl 1991). Moreover,
holes attributed to emergence of S. granarius were
found in charred barley grains in a cesspit deposit
of the early 14th century AD in Svendborg, Denmark
(Jørgensen 1986), although, as mentioned earlier,
“bubbling” during charring can produce holes strongly
resembling those caused by weevils. Remains
named as S. oryzae were identified from the wreck of
the “Amsterdam” by Hakbijl (1987). King (unpublished)
found S. granarius in 13th-century latrine deposits
in Riga, Latvia. Deposits dated to the 14th century
in Novgorod, Russia, yielded equal, though very
small, numbers of S. granarius and O. surinamensis.
Ptinus villiger (Reitter) was also recovered from the
site (Hellqvist 1999).
In Ireland, the period following the Norman
Conquest heralded the first appearance of the classic
grain beetles. The earliest fossil record of the
granary weevil in Ireland is from late Viking/early
Anglo-Norman deposits at Waterford (Reilly 1994).
The scavenger B. lethifera, present in Viking Age
deposits, was found in late 12th-century layers at
Essex Street West, Dublin, and S. granarius was recovered
from early 13th-century deposits at the same
site (Reilly 2003). The granary weevil has also been
recovered from late 13th/early 14th-century layers of
2014 Journal of the North Atlantic No. 23
G.A. King, H. Kenward, E. Schmidt, and D. Smith
10
and often protected within grains. In experiments,
a range of insects survived charring in good condition
up to moderate temperatures, though they were
severely damaged above around 400 °C (Kenward et
al. 2008). Nevertheless, while the charred insects appear
to be in superb condition, with appendages and
even hairs and setae entire, they are very fragile and
easily broken into fragments small enough to evade
recovery. Are we even losing the tiny sclerites of
waterlogged Cryptolestes through our sieves? There
may be problems of this kind; nevertheless we can
certainly piece together a worthwhile story, albeit
with many gaps.
What was the infested grain intended for?
What do the records of grain pests tell us about the
past? It may seem obvious that they were important
in damaging stored grain intended for people.
But did they appreciably reduce the availability of
food? Hall and Kenward (1976) suggested that the
structure at Coney Street in York mentioned above
may have been a transit shed and thus a serious
source of infection for the grain passing through it,
leading to contamination in the main granary. This
explanation now seems far less certain in view of
our understanding of the overwhelming presence of
stable manure in Roman towns and military centers,
with grain pests being part of the characteristic “indicator
group” for this material (Kenward and Hall
1997). For Roman and Post-Conquest Britain, many,
if not most, records of grain beetles now seem to be
from horse feed deposited as stable manure (Kenward
2009; Kenward and Hall 1997; King, in press
c; Smith and Kenward 2011). We are left to wonder
whether horse feed was specially stored, with little
regard to infestation, or whether perhaps grain originally
intended as human food was routinely diverted
to equines if it became infested. If so, the abundant
grain pests would be of far less significance in our
reconstructions of past human life; this question and
the wider issue of the routes by which grain pests
entered deposits, have been discussed by Smith and
Kenward (2011, 2012).
Fortunately, not all of the fossil grain pest assemblages
seem to have been from animal feed, as
those from a number of Roman and Post-Conquest
deposits lack key components of Kenward and
Hall’s (1997) stable-manure indicator group. For
example, the grain pests recovered from the Invereskegate
well (Smith 2004) are more indicative
of grain dumping rather than conversion to fodder.
Moreover, the infested cereals may not have
always been intended for consumption. Analysis
of the modern insect fauna from The Oldest House
at West Stow, Sussex suggests that grain pests may
(F.), and both S. granarius and S. oryzae (Bain
1998, Bain and King 2011). Deposits dating from
the 18th century in Québec City yielded S. granarius,
Tenebrio molitor, Alphitobius bifasciatus, and
Ptinus fur (Bain et al. 2009). Also, Bain (2001)
recorded Rhyzopertha dominica, Sitophilus oryzae,
S. granarius, Cryptolestes, Tribolium castaneum, T.
confusum, Tenebrio molitor, and T. obscurus from
deposits from Québec City dated to the 19th century,
in assemblages dominated by Old World species. In
early to mid-19th-century deposits from Toronto in
Ontario, Canada, cf. Blaps sp., Ptinus cf. clavipes,
P. fur, Sitophilus granarius, and O. surinamensis/
mercator were recovered alongside the house fauna
beetles Mycetaea subterranea and Typhaea stercorea
(King 2010a). Moreover, by the 18th century,
certain grain species had arrived on the west coast of
North America; Essig (1927) reported evidence of S.
granarius and S. oryzae in abode bricks of Santo Domingo
Mission in lower California, dated to 1775.
The spread of storage pests to the rest of the
world
Analyses of archaeological fossil insects are
lacking for other lands colonized by Europeans,
but early collections and published records show
the arrival of the storage pests with colonists, and
subsequently with trade—hardly surprising as the
majority are now cosmopolitan. Many alien insects
are certainly established in Australasia, especially
in New Zealand (e.g., Cumber 1961, Kuschel 1979,
Moeed 1993). Archaeological investigation in Australia
and New Zealand may well produce evidence
of the early arrival of the grain pests, and also of the
various other insects that have entered the ecology
of those countries. The situation is similar for South
America: archaeological investigation of colonial
period sites is awaited, but worthwhile results can
surely be anticipated.
Discussion and Future Directions
How good is the fossil record?
We have already issued a caveat concerning
corruption of the evidence by contaminants and
misidentifications. There may also be a taphonomic
(preservational) effect. The different potential for
preservation in different regions has already been
alluded to. In addition, the more delicate taxa are
potentially under-represented. In general, this probably
does not undermine the record since storage
insects preserved by anoxic waterlogging, desiccation,
and charring all contribute. There may be some
bias, however. Charring may favor Sitophilus over
the other common grain pests because it is tough
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G.A. King, H. Kenward, E. Schmidt, and D. Smith
A current limitation of the archaeobiogeographical
approach is that it is unable to differentiate
between multiple introductions of a species, as it
does not distinguish between different populations
of the same species. The Roman garrisons appeared
to have been receiving cereal supplies from several
provinces during the early Roman conquest and occupation
of Britain. As populations of Sitophilus
granarius presumably existed in stores around the
Mediterranean and further north, the species was
probably transported to Britain from multiple regions
on many occasions. However, an archaeobiogeographical
study of S. granarius would be blind to
these introductions, so that details of trade connections
to regions with early Roman settlements would
be obscured. Thus, in areas such as 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, the records give only the crudest
view of culture contact. However, the recovery
of other grain fauna might provide further insight
towards any potential trade connections, which may
enhance the efficacy of the archaeobiogeographical
approach but which would perhaps also risk placing
undue confidence or faith in the currently patchy fossil
accounts of species’ distribution.
Closer analysis of the fossils might be rewarding.
The suspected “contaminants” from the early
medieval era in Britain mentioned above might be
AMS dated to determine if they are redeposited or
intrusive, and thus if pests survived or were only
re-introduced after the Norman Conquest. Analyses
of genetic diversity in fossils in the Roman and Norman
period onwards might give clues. DNA has been
recovered from Roman and later fossils of Sitophilus
from English sites (King et al. 2009), suggesting the
approach might be feasible. Are Roman and Norman
populations genetically distinct? Investigating
this would necessitate the recovery of abundant
exceptionally well-preserved fossils from numerous
sites. In many areas, and probably in most towns,
the buried organic heritage is under threat (Kenward
and Hall 2008), and needs to be protected so that the
available stock of such remains does not diminish
before it can be examined.
Regarding the status of the pests in towns and
military establishments—whether they were depleting
resources for human consumption or exploiting
low-grade cereals primarily intended for
livestock—we might examine the relationship of the
grain community to decomposer and domestic fauna
in deposits where preservation is good and insects
abundant. Such analysis at some British sites has
enter along with stored, unprocessed cereals used as
thatch (King 2010b, 2012). Archaeologically, this
may offer a possible explanation for the presence
of grain pests in Smith et al.’s (1999) late medieval
thatch from Southern England, and similarly on
the other side of the Atlantic, in the colonial fort at
Jamestown, Virginia (King et al. 2010).
Another significant route by which grain pests,
and particularly Sitophilus, appears to have entered
archaeological deposits is in human feces, for it
has been shown that grain pests can pass relatively
undamaged through the human gut (Osborne 1983).
Certainly, grain pests are common enough in cesspits
(cf. Smith 2013).
Efficacy and reliability of the archaeobiogeographical
approach
When married to the archaeology, biogeography
can serve as an effective tool for investigating a
wealth of past human activities. By studying the fossil
record for distributional changes in grain fauna
over time, a number of archaeologically significant
themes, such as past patterns of human migration,
cultural contact, and trade, can be elucidated, for
example. Unfortunately, an archaeobiogeographical
approach also suffers from limitations, with the reliability
of results hedged by the quality of the fossil
record.
The study of insect remains from archaeological
contexts is still rarely carried out outside of the British
Isles, an issue addressed by Buckland (1981). The
resulting lack of data continues to pose problems,
particularly in attempting to ascertain shifts in species’
distributional ranges. Thus, although Sitophilus
granarius has been recovered from early Holocene
occupation sites in central Europe, the next record
of the species in northern or central Europe does not
occur until the Roman occupation. Was the granary
weevil absent from the region during the Bronze and
Iron Ages? Or did central Europe remain a potential
source for the diffusion of the species into later prehistoric
sites in the rest of Europe? It is only when
numerous archaeological analyses have been made
that the species’ absence from the region between
the Neolithic and Roman Periods can be inferred.
For example, trade and cultural contact between
Britain and mainland Europe during the Bronze and
Iron Ages would potentially have provided opportunities
for the introduction of extant populations of
the granary weevil to Britain prior to the Roman Era,
were it at all common at mainland sites. Certainly,
the grain pests seem to have spread easily enough in
the Mediterranean region, and much more recently
around the North Atlantic.
2014 Journal of the North Atlantic No. 23
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12
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proved informative, sometimes strongly suggesting
that they indeed originated in stable manure,
and occasionally hinting that they came from some
other source (Kenward and Carrott 2006, Smith and
Kenward 2012). Similarly, the application of stable
isotope analyses to insect fossils in parallel with the
remains of domestic vertebrate fauna may prove
useful as a means of approaching this question (see
King, in press a).
In conclusion, we have a very substantial fossil
record for a range of grain beetles, but it is very
patchy and predominantly from northwest Europe;
no area yet has an adequate record. If we are to understand
their origin and dispersal, fossils are needed
from across the Palaearctic and North Africa. The
analysis of pre-agricultural natural deposits in the
Middle East, North Africa, and the Indian Subcontinent
might greatly advance our understanding of the
original ranges of the pest species and the stages by
which they entered into association with humans.
And finally, further research along these lines could
address the question: have any of the storage pest
species, and especially Sitophilus granarius, actually
evolved in response to human activity?
Acknowledgments
G.A. King’s research was supported, in part, by the
Short-term Marie Curie PALAEO Fellowship at the University
of York, UK, and a Junior Research Fellowship
at the Durham University. H. Kenward is grateful to the
University of York’s Department of Archaeology for postretirement
access to facilities. We also wish to extend
our thanks to Jamestown Rediscovery and Preservation
Virginia for their contributions.
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