Journal of Archaeological Science 1975, 2,85-94

Pitfalls in the Environmental Interpretation of Insect Death Assemblages Harry K. Kenward Unit for Environmental Research in Archaeology, York Archaeological Trust, York YOl 2BX, U.K.

Attention is drawn to the absence of an established basis for the reconstruction of past environments from insect death assemblages. A naive reconstruction based on a modern death assemblage from a drain in the City of York is tested and shown to be inadequate in a number of respects. Certain of the habitats indicated by the fauna are absent from the surroundings, while the importance of some others is exaggerated. It is concluded that these errors were brought about by the active or passive long distance transport of insects.

Introduction A considerable number of reconstructions of past ecological conditions based on archaeological insect assemblages have been made (for access to literature see Kenward, 1974). The ways in which, it is thought, the archaeologist might profit from such work have been outlined on several occasions (Osborne, 1973; Speight, 1974; Kenward, 1974) but there has been no detailed consideration of the theory behind, or test of the accuracy of, these interpretations. Discussion of the use of insect faunas in archaeology by Speight (1970, 1974) offers only the suggestion that “indicator families” should be used to reduce the work load in identification and that keys for the identification of insects from individual sclerites must be constructed. These views are not supported by other workers. I have used an account of the literature and archaeological records of the beetle Aglenus brunneus (Gyll.) to illustrate the difficulty of determining the implications of a single species in archaeological assemblages (Kenward, 1975). The use of a large number of species and individuals in making reconstructions is urged and it is suggested that species associations characteristic of particular habitats must be recognized.

I have, over several years, studied a variety of modern insect death assemblages to obtain an understanding of their formation. In the present paper the reliability of a simple reconstrtiction from a death assemblage formed in an urban situation is tested by comparison with the recorded ecology of its surroundings.

Methods The insect remains were extracted from a deposit formed in a drain sump in an alleyway behind Fern Street, York (Grid Reference SE 606 524). The sample was scooped from the sump with a ladle and by hand, carried to the laboratory in a covered container and

85

86 H. K. KENWARD

processed immediately. After washing and paraffin separation (see Kenward, 1974) the concentrate was sorted under the binocular microscope and remains of Coleoptera, Hemiptera-Heteroptera and some other insects picked out. These were identified by comparison with named material and the number of specimens of each taxon represented by them computed. The known ecology of each species was then tabulated and the data collated to form a description of the surroundings of the deposit. For this purpose the naYve assumption was made that the breeding habitat for each of the species must have been present near the point of deposition.

Results The material was collected on 27 March 1974. The sample, whose volume was about 2500 cc and weight about 3 kg, consisted of a foul-smelling, black, reduced slurry containing a great deal of water. It included some small stones, sand and finer mineral particles, twigs, leaves, seeds and a variety of small artifacts.

The specimens identified included a few almost complete individuals but were mostly recognized from isolated sclerites. The minimum number of individuals of each taxon of Coleoptera and Heteroptera which could have given rise to the recorded sclerites are listed in Appendix 1. A total of 259 specimens of 115 species are represented. The specimens are sorted in spirit or on card slides in the author’s collection and are labelled with the code “MOD 8”.

The Implications of the Fauna A short summary of the habitats of each of the recorded species is given in Appendix 1 (Table Al). The greater part of this information is taken from Donisthorpe (1939), Fowler (1887-1891), Hoffmann (1950-1958), Lindroth (1974), Southwood and Leston (1959) and is supplemented by the author’s unpublished data. Other sources are given where appropriate in the Table. These summaries have been used to construct Table A2, in which the number of species and individuals are given for each of a series of broad habitat classes. This process is regarded as representing in tabular form the reasoning which has been employed in interpreting archaeological faunas. The true range of habitats of each stage of the life history of most species is poorly known; sources offer different information, while collectors’ impressions of the main habitats of a species vary. This has made the choice of species placed into each of the sections of Table A2 somewhat arbitrary, but not in a way likely to have influenced the results. Many species fall into two or more of the habitat classes.

The implications of the fauna using the method of summation of habitats are briefly as follows.

The presence of several aquatics suggests that the deposit formed in an open, water- filled depression. The surroundings were open ground with a short vegetation cover which probably included weeds of disturbed ground, including nettles. Some bare soil was, however, present. There was abundant dead wood close to the point of deposition, as well as the dung of herbivores and large volumes of rotting plant matter. The influence of man was very strong and buildings, domestic or for storage, were very close and probably in use.

The surroundings of the drain may be summarized as follows. The locality lies in an area of terraced houses in a completely built up area. The deposit formed in a drain sump of narrow aperture in a concreted alleyway between high brick walls. The surroundings of the drain are barren, at least 95 % of the area being covered by buildings, roads or paved paths and yards, over a radius of 250 m. There are small areas of bare earth in

PITFALLS IN INSECT DEATH ASSEMBLAGES

some yards, occasionally with scattered grasses and weeds, but in insignificant amounts and not close to the drain. There is no open water which could provide habitat for, or attract, aquatic species. The nearest open water is probably in unrecorded garden ponds, but not within 100 m. Nettles may be present but have not been seen. Wood, both seasoned and rotten, is present. There is no dung of herbivores anywhere nearby. Rotting vegetation in accumulations suitable for a compost fauna is not recorded.

Discussion Deposits formed in depressions which would have received run off water are often encountered during archaeological excavations. Few sediments are laid down in situations where there is no access to the open air.

The modern deposit discussed here is valuable in that it has no autochthonous fauna or species breeding in large numbers immediately adjacent. This means that no species should be over represented through proximity of its habitat. There is no reason to suppose that, in general terms, this deposit cannot be compared with any deposit formed in a hollow into which water drained from the ground or from roofs, for example an un- protected well, pit, ditch or merely a damp yard or rubbish heap.

The sediment had almost certainly accumulated during the year or two preceeding its collection as no insect remains showed a high degree of decay. The sump is liable to partial flushing by rain. It is considered that the silt contained a reasonably good random sample of the insects available to it, but that only a small proportion of the individuals entering the drain have been retained in the sump, the remainder having been carried away by the flow of water. The drain receives water from an estimated 12 m2 of paths.

The summation of habitats provides a description of the surroundings which is in many respects quantitatively and qualitatively inaccurate. The number of insects placed in each habitat class does not reflect the relative abundance of the habitats in a 250-m radius. Decaying masses of vegetation and open ground with a cover of short vegetation are over represented, while two habitats, open water and the dung of herbivores, are indicated but absent in the vicinity. Had this been a fauna from an archaeological site the inferences drawn from it would have been incorrect. A deposit which received run- off from a well-kept courtyard, for example, might give a large fauna suggesting squalor and decay.

There are two reasons why this should be so. First, it is likely that some of the species can breed in pieces of habitat which appear to be insignificant, for example, small plant tufts and detritus-filled cracks. Such situations are rarely examined by collectors and so rather more species than recorded may be able to breed in even the most barren “urban desert”. A large number of insects can, of course, breed in towns and will contribute to the “background fauna” discussed below, but this does not detract from the present study, in which a very restricted and exceptionally barren area is being considered. Only a propor- tion of the species may be accounted for by local breeding even if the surroundings up to a 250-m radius are considered. In archaeological investigations information about much smaller areas may be required, perhaps up to a radius of only a few metres from the point of deposition; often the autochthonous fauna alone is of interest.

The second, and in the present context more significant, reason for the occurrence of this large and diverse fauna is the habit of dispersive flight at some stage during the adult life of most insect species. Only five of the beetles recorded (Pterostichus madidus, Anommatus duodecimstriatus, Metophthaima serripennis, Otiorrhynchus sulcatus and Liosoma de$exum) are invariably flightless. Many of the remaining species are ready flyers and almost all have been recorded on the wing or in circumstances suggesting flight. In the present case some of the insects probably covered a considerable distance

88 H. K. KENWARD

before settling in the drainage area of the sump. The local prevailing wind, which would determine the flight direction of all but the largest species, is channelled across a large tarmac car park from the barren centre of the city.

Almost all the Coleoptera of the present fauna have been found in death assemblages formed in roof gutters. The only exceptions are the rare Metophthalma serripennis and the uncommon and cryptic Anommatus duodecimstriatus. These roof assemblages, in which nearly 300 species of Coleoptera have already been found, must have formed as a result of flight, wind transport, phoresey or the accumulation of bird droppings. This work, the literature of flight in the British Coleoptera and Heteroptera and the importance of this data in the interpretation of ancient faunas, will be reported elsewhere (in prepara- tion). The composition of many of the archaeological faunas examined by the author could be largely accounted for by this “background fauna”. Its importance will be exaggerated when population densities are low at the point of deposition and where run off is received, as in the present case and in many situations encountered in archaeology.

Conclusions This work has shown that a simplistic approach to the interpretation of archaeological faunas, using the summation of the habitats of the recorded species alone, is not always reliable. It is probable that many insects fly, or are passively transported, over con- siderable distances before becoming incorporated in a deposit and that in some circum- stances a large proportion of a death assemblage may originate in such a way. An understanding of the composition of this background fauna is vital.

These results are relevant to the study of faunas of “natural” deposits, including those of Pleistocene age, as well as of anthropogenic sediments. Where the regional ecology and climate are being investigated the phenomenon of long distance transport means that at least a proportion of the species living in a wide area will be represented. The ability to distinguish the regional component of a death assemblage on the one hand and the local component on the other is essential if palaeoentomology is to be a reliable tool for the accurate investigation of the past.

Appendix 1 Table Al. Records of Coleoptera and Hemiptera from drain sump, Fern Street, York

Taxon

Carabidae Nebria brevicollis (F.) Notiophilus biguttatus (F.) Clivina fossor (L.) Bembidion Iampros (Hbst.) B. quinquestriatum Gyll. B. ?guttlrla (F.) Trechus quadristriatus (Schr.)

Bradycellus verbasci (Duft.)

Amara sp.

Pterostichus madidus (F.)

Minimum number Recorded ecology

Coleoptera

1 Eurytopic ground beetle. 1 On somewhat shady open ground, usually in dry places. 1 All kinds of dampish, usually vegetated, open ground. 2 Dry, open soil; eurytopic. 3 Synanthropic, in dry places. 3 Open ground, shaded or near water. 1 Open rather dry places with short vegetation. Damp

places (Joy, 1932). In compost. 1 On sand or gravel soil amongst low plants. Comes to

light. 1 All more or less xerophilous. on open ground with short

plants. 2 Open country, often cultivated soil, in gardens.

PITFALLS IN INSECT DEATH ASSEMBLAGES

Table Al continued

Taxon Minimum number Recorded ecology

Dytiscidae Agabus bipustulatus (L.)

Hydrophilidae Helophorus grandis (Ill.)

(aquaticus auctt.) H. cf brevipalpis Bed.

Sphaeridiidae Cercyon haemorrhoidalis (F.) C. unipunctatus (L.) C. spp. indet. Megasternum obscurum (Msh.)

Clambidae Clambus pubescens Redt.

Staphylinidae Micropeplus fulvus Er. Proteinus ?ovalis St.

Omalium rivulare (Payk.)

Xylodromus concinnus (Msh.) Omaliinae indet. (not above) Oxytelus rugosus (F.) 0. laqueatus (Msh.) 0. sculptus Grav. 0. sculpturatus Grav. 0. nitidulus Grav. 0. complantatus Er. 0. tetracarinatus (Block) Platystethus arenarius (Geoff.) Stenus spp. indet. Lathrobium sp. indet. Xantholinus ?angustatus St. X. linearis (01.) X. longiventris (Heer.) Neobisnius ?cerrutii Philonthus sp. indet. Gabrius sp. indet. Quedius sp. indet.

T. hypnorum (F.) or chrysomelinus (L.) T. spp. indet. (not above)

Tachinus rujipes (Deg.) T. ?laticoilis Grav. ?Aleochara sp.

Aleocharinae indet. spp.

Pselaphidae Tychus niger (Payk.)

Tachyporus ?obtusus (L.)

1 Almost all kinds of still or sluggish fresh water (Balfour- Browne, 1950).

1 Still or sluggish fresh water; eurytopic (Balfour-Browne, 1958).

5 As H. grandis.

1 + ?I Compost, dung, old bones. 1

10

3 2

2

1

1 3

1

4

1

1

1 5

1

1

16

1 1 1

31

1

5 1 2 1 1 2 2 1 2 1 1

1

Dung, wet rotting vegetation, corpses. Decaying plant and animal matter, dung, in moss, etc. Rotting plant matter; at roots of plants.

Decaying plant matter; probably mould feeder (Johnson, 1966).

Rotting plant matter; in soil (Tottenham, 1954). Rotting plant and animal matter (Tottenham, 1954); dung. Decaying plant and animal matter, dung, nests, fungi, moss. Haystacks and vegetable refuse; fungi, nests. Eurytopic as subfamily. Rotting plant matter, dung; mud by water. Dung, rotten fungi. Rotting plant matter. Rotting plant matter. Rotting plant matter; mud by water. Rotting plant matter Rotting plant and animal matter; dung. Dung, rotten fungi. Eurytopic as genus. In decaying matter, roots of plants, under bark etc. In moss and vegetable refuse. Roots of plants; in moss, compost, leaf litter. Plant refuse; at roots of plants. Rotting plant matter (Horion, 1965). Plant roots, compost, dung, corpses etc. Decaying plant matter; at roots of plants, in moss etc. Decaying plant and animal matter; in moss, at roots of plants etc. At roots of plants, in moss, compost etc. At roots of plants, in compost, moss, nests etc. Dead plant matter; at plant roots, in moss, on blossom etc. At plant roots, in decaying matter, dung etc. At roots of plants, in compost. At roots of plants, in decaying matter, dung etc. Almost ubiquitous.

In decaying plant matter, moss etc.

90 H. K. KENWARD

Table Al continued

Taxon Minimum

number Recorded ecology

Dermestidae Anthrenus fuscus 01.

Nitidulidae Meligethes sp. indet. Omosita discoidea (F.)

Rhizophagidae Rhizophagus perforatus Er.

Cryptophagidae Cryptophagus spp. indet. (at least

two species) Atomaria spp. indet. (at least two

species)

Lathridiidae Stephostethus Iardarius (Deg.)

Aridius nodifev (Westw.) A. bifasciatus (Reitt.) Enicmus spp. indet. (at least two

species) Metophthalmus serripennis (Broun.)

Corticaria, Corticarina or Melanophthalma spp.

Mycetophagidae Typhaea stercoraria (L.)

Colydiidae Anommatus duodecimstriatus (Miil.)

Endomychidae Mycetaea hirta (Msh.)

Coccinellidae Adalia bipunctata (L.) A. decimpunctata (L.) Coccinella septempunctata L.

Anobiidae Anobium punctatum (Deg.)

Ptinidae Ptinus tectus Boield.

Anthicidae Anthicus quisquillius Thorn. or

or jloralis (L.)

Mordellidae Anaspis costai Emery

1 Reported to feed on dead insects in spider webs (Hinton, 1945). Adults on flowers.

2 Phytophagous, wide variety of hosts; adults on blossom. 2 Rotting plant and animal matter, bones; on blossom.

1 In rotting plant matter; under bark; on fungi.

6

4

In dead plant matter, at roots of plants; under bark; in fungi etc. As Cryptophagus.

1

4 2 4

3

14

3

3

5

3 1 1

4

2

1

1

In compost, at roots of plants, in moss, on dead twigs etc. Decaying plant matter; fungi; at roots of plants etc. Compost, at plant roots; often in urban situations. Compost, moss, roots of plants, under bark etc.

Synanthropic but perhaps becoming established out of doors in compost, straw etc. Decaying plant matter, straw, at plant roots etc.

Rather dry decaying plant matter, nests; stored products (Hinton, 1945).

Vegetable refuse (Joy, 1932); decayed seed potatoes and wood underground.

Decaying plant matter; fungi; in damp buildings (Hinton, 1945).

On a wide range of herbs, shrubs and trees; predatory. As A. bipunctata. As A. bipunctata.

Dead wood of many kinds.

In buildings; nests.

Both in rotting plant matter (Buck, 1954).

Larvae in wood (Buck, 1954); adults on blossom.

PITFALLS IN INSECT DEATH ASSEMBLAGES

Table Al continued

91

Taxon Minimum number Recorded ecology

Scarabaeidae Aphodius rujipes (L.)

A. prodromus (Brahm)

Chrysomelidae Longitarsus sp. indet. Haltica sp. indet. Chaetocnemu concinna (Msh.)

Curculionidae Apion sp. indet. Otiorrhynchus sulcatus (F.) Phyllobius pyri (L.) Sitona hispidulus (F.)

Sitonu spp. indet. (not including above)

Pentarthrum huttoni Woll. Euophryum confine (Broun) Liosoma ?defie.wm (Pz.)

Cidnorhinus quadrimaculatus (L.) Ceuthorhynchus erysimi (F.) C. contractus (Msh.) C. sp. indet. (not above)

Lygaeidae Drymus sylvaticus (F.)

Cimicidae Anthocoris ?nemorum (L.) O&s sp. indet.

Miridae Liocoris tripustulatus (F.)

Lygus sp. indet.

Saldidae Saldula saltatoria (L.)

Corixidae Sigara sp. indet.

1

1

2 1 4

1 1 1 1

5

2 15

1

1 1 5 1

Dung of herbivores; v. rarely in dung-like wet rotten plant matter. Dung of all kinds; rotting plant matter (Landin, 1961).

Phytophagous; on a wide range of herbs. Phytophagous; various herbs. Phytophagous; various herbs; especially Polygonum (Joy, 1932).

Phytophagous; various herbs and shrubs. At the roots of plants. Phytophagous; various Rosaceae; nettles. Phytophagous; various Papilionaceae, especially Tri- folium. Phytophagous; mainly Papilionaceae.

Decayed wood. Decayed wood. Phytophagous; various Ranunculaceae, especially Ranunculus, Anemone. Urtica, Parietaria (Davis & Lawrence, 1974). Phytophagous; various Cruciferae. As C. erysimi. Phytophagous; wide host range.

Hemiptera-Heteroptera

1 At roots of low vegetation.

6 Predatory; on a wide range of herbs, shrubs and trees. 1 As A. nemorum.

1 Phytophagous; Urtica, Parietaria (Davis & Lawrence, 1974).

1 Phytophagous; wide host range.

1 Damp open ground by water; exceptionally in drier places (Kenward, 1970).

1 Aquatic; most kinds of still and sluggish water.

Table A2. Number of species and specimens in each of several habitat classes

Ecological group Number of

species specimens Implications Actual ecology

Aquatic True aquatics At water margin

4 8 Presence of this group sug- Not a water body of this 1 1 gests a deposit formed in kind-sump water would not

quiet water, e.g. ditch, pond, support or attract these water-filled pit. species.

92 H. K. KENWARD

Table A2 continued Number of

Ecological group species specimens Implications Actual ecology On open ground 7 12

At roots of low plants (i.e. often found amongst the matted stems and detritus at the base of herbs)

Phytophagous True phytophages

Secondarily tied to plants by predation In rotting plant matter Mostly in this habitat Able to exploit the habitat Ditto, generically named

In dung Primarily in (herbi- vore) dung

Able to exploit dung 8 21

In dead wood exclusively

Able to exploit rotten wood

With man Strongly synanthro- pit Often in direct association with man, in buildings Ditto, generically named Often found in areas disturbed by man

24 34

15 28

5 12

8 38 27 58

6 14

4 4

4 22

2 8

2 6 18 57

8 29

whole

Surrounding of open or sparsely vegetated ground ; probably fairly dry, sunny, damper and shaded in places.

It is not feasible to divide this group into “mainly” and “sometimes” found in this situation. None are confined to it but this group, with the following one, suggests a good cover of low plants in places.

Indicate Ranunculaceae, Urticaceae, Cruciferae, Papilionaceae (probably dis- turbed ground weed flora) and Rosaceous shrubs.

Indicate only the presence of living plants.

Abundance of species of de- caying plant matter and those facultative to it indicates the presence of masses of com- post-like material. Possibly the deposit included some of this matter. Such habitats are man-made.

All have been recorded in wet rotting vegetation (one only very exceptionally) but as a group suggest presence of herbivore dung. Lend weight to the above evidence but all also found in rotten plant matter.

Strong indication of dry seasoned and slightly damp rotten wood near-by. Mouldering dead wood but both also able to utilize compost.

Taken as a group strongly suggest the area to be greatly influenced by the activities of man, and that buildings may be present. All the species have, however, been found in the wild in some circum- stances. The whole fauna falls in this category, even the aquatics being character- istic of garden ponds.

Small patches of habitat for some of these in yards but not in alleyway.

No extensive short plant community, only isolated plants.

Plants of these groups not at all abundant. Very small numbers of Cupsella (Cruci- ferae) in yards but no other members of these families recorded. Some rose bushes. A few suitable plants for these in yards.

Although small amounts of dead plant matter present, it is not abundant and none is close to the sump.

No dung of herbivores (or suitable wet rotting vege- table matter) present.

Moderate amounts of dry timber and mouldering wood within a few metres.

No natural habitats present in the entirely man-made landscape.

PITFALLS IN INSECT DEATH ASSEMBLAGES 93

Appendix 2

A note on the seed remains

By Dorian Williams Unit for Environmental Research in Archaeology, York Archaeological Trust.

The seeds extracted from the drain sump deposit by paraffin flotation have been examined. The species identified are listed in Table Bl.

The relationship between deposited seeds and the surrounding environment was not easy to test from this sample, due to the nature of the site and the restricted range of seeds recovered. The species are also unsatisfactory as environmental indicators.

Poa unnua is the only common species likely to be found locally, in garden plots, lawns and even crevices in pavements. A few small tufts of this species were growing at the base of the alley walls. The other two dominant species, and probably Setariu itafica, were derived from a commercial bird seed mixture. None of these species would have achieved such prominence in an archaeological deposit, as a result of the poor preservation of Gramineae seeds and their consequent difficulty of identification.

The presence of tree species is not unexpected as their seeds are wind dispersed. However, the relatively large numbers found are surprising in view of the absence of trees in the immediate vicinity. On an archaeological site this might have led to a mis- interpretation of the evidence, although prior knowledge of the urban status of the site may have led to some caution.

The presence of an open, disturbed environment is indicated to a certain degree by the weed species. Sufficient small habitats exist in the vicinity to accommodate these, although few of them have been recorded. In view of the limitation of the grass and tree species discussed above, archaeological interpretation would have been based on these ruderal species. The small amount of evidence they provide can be considered to reflect fairly closely the sterile nature of the surroundings.

Table Bl. The seed remains recorded from the Fern Street drain sump

Species Number of specimens

Poa annua L. Panicum SQ. cf P. mileaceum L. Phalaris SQ. cf P. canariensis L. Betula sp. cf B. pendula Roth. Fraxinus excelsior L. Setaria sp. cf S. italica (L.) Beauv. Chmaenerion angustifolium (L.) SCOQ. Rumex SQ. Chenopodium rubrum L. Polygonum aviculare L. Urtica urens L. Achiliea millefolium L. Taraxacum sp.

f%WUdS SQ.

Compositae

These grasses were the dominant species. Each gave counts well in excess of 100. 50+ 20 2

1

1 Several fragments of stone.

Unattached pappus.

Acknowledgements

I am grateful to Janet Leng, who showed me the study site and has helped me in various other ways. Tish Farber, Richard Hall, Richard Morris, Peter Osborne, Ben Whitwell and Doria, Williams have each given me their helpful advice during the preparation of the manuscript.

H. K. KENWARD

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