The Bugs Coleopteran Ecology Package (BugsCEP) database: 1000 sites and half a million fossils later

Quaternary International xxx (2014) 1e11

Contents lists available at ScienceDirect

Quaternary International journal homepage: www.elsevier.com/locate/quaint

The Bugs Coleopteran Ecology Package (BugsCEP) database: 1000 sites and half a million fossils later Philip I. Buckland Department of Historical, Philosophical and Religious Studies, Environmental Archaeology Lab, Umeå University, Umeå SE-90187, Sweden

a r t i c l e i n f o

a b s t r a c t

Article history: Available online xxx

The Bugs database project started in the late 1980s as what would now be considered a relatively simple system, albeit advanced for its time, linking fossil beetle species lists to modern habitat and distribution information. Since then, Bugs has grown into a complex database of fossils records, habitat and distribution data, dating and climate reference data wrapped into an advanced software analysis package. At the time of writing, the database contains raw data and metadata for 1124 sites, and Russell Coope directly contributed to the analysis of over 154 (14%) of them, some 98790 identifications published in 231 publications. Such quantifications are infeasible without databases, and the analytical power of combining a database of modern and fossil insects with analysis tools is potentially immense for numerous areas of science ranging from conservation to Quaternary geology. BugsCEP, The Bugs Coleopteran Ecology Package, is the latest incarnation of the Bugs database project. Released in 2007, the database is continually added too and is available for free download from http:// www.bugscep.com. The software tools include quantitative habitat reconstruction and visualisation, correlation matrices, MCR climate reconstruction, searching by habitat and retrieving, among other things, a list of taxa known from the selected habitat types. It also provides a system for entering, storing and managing palaeoentomological data as well as a number of expert system like reporting facilities. Work is underway to create an online version of BugsCEP, implemented through the Strategic Environmental Archaeology Database (SEAD) project (http://www.sead.se). The aim is to provide more direct access to the latest data, a community orientated updating system, and integration with other proxy data. Eventually, the tools available in the offline BugsCEP will be duplicated and Bugs will be entirely in the web. This paper summarises aspects of the current scope, capabilities and applications of the BugsCEP database and software, with special reference to and quantifications of the contributions of Russell Coope to the field of palaeoentomology as represented in the database. The paper also serves to illustrate the potential for the use of BugsCEP in biographical studies, and discusses some of the issues relating to the use of large scale sources of quantitative data. All datasets used in this article are available through the current version of BugsCEP available at http:// www.bugscep.com. Ó 2014 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction

searches when looking for information on the habitat of beetles found fossil. Perhaps most usefully, it provided a means to rapidly summarise the habitat information, bibliographic data and known fossil record of all taxa found at a site. Environmental interpretations and reconstructions could thus be more efficiently undertaken. From the very start the aim was to partner advanced database tools and interfaces with the database, rather than simply making data available, and later versions have continued to be developed in this spirit. Subsequently, the system has been through various revisions and improvements, with the latest version becoming an advanced

The Bugs database was initiated in the 1980s as a database of coleopteran fossil records and ecological information to be used as a tool in interpreting Quaternary entomological data (Sadler et al., 1992). It was used to collate data and information from disparate sources in the international entomological literature and analogue fossil insect datasets, reducing the need for extensive library

E-mail addresses: [email protected], [email protected]. 1040-6182/$ e see front matter Ó 2014 Elsevier Ltd and INQUA. All rights reserved. http://dx.doi.org/10.1016/j.quaint.2014.01.030

Please cite this article in press as: Buckland, P.I., The Bugs Coleopteran Ecology Package (BugsCEP) database: 1000 sites and half a million fossils later, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2014.01.030

2

P.I. Buckland / Quaternary International xxx (2014) 1e11

software application for aiding studies in, to name but a few, Quaternary science, archaeology, ecology and conservation (Buckland and Buckland, 2006; Buckland, 2007). The database is also extensively expanded, both in terms of scientific scope and the amount of data. The work of Russell Coope and colleagues has directly contributed to a significant part of the data held in Bugs, and it could be argued that Russell is indirectly responsible for almost the entirety of the rest of the data by way of his former students and colleagues. 2. System origins, evolution and usage The Bugs system began as a DBase4 database coupled with interface software developed in Clipper (Sadler et al., 1992). The evolution of Bugs from this version to the current one is described by Buckland (2007), and the present iteration includes a considerable expansion in terms of scope and application. Although originally designed as a tool for archaeology, palaeoecology and the Quaternary sciences, Bugs has potential applications far beyond these fields. The full potential, especially with respect to its use in modern ecology, biogeography and conservation sciences, has yet to be realised, but the list of known publications using or referring to the system indicates an expansion beyond the initial target user group (Fig. 1). To this date, at least 100 publications have acknowledged the use of Bugs or BugsCEP as a source of data, data repository or research tool since 1997 (see http://bugscep.com/publications.html). Of the 93 publications citing the database and identified on the internet by the authors, 58% of them could be considered as being outside of the initial project scope. (The authors would be most grateful for any additions to this list, irrespective of whether the publication cites the database or not). The data in BugsCEP are being continually updated, and new software and accessibility tools are under development within the scope of the SEAD (Strategic Environmental Archaeology Database) project (http://www.sead.se; Buckland, 2014; Buckland and Eriksson, 2014; Buckland et al., 2010). The statistics and enumerations presented in this paper reflect the status of the Bugs database on the 17th June 2013. 3. Geographical, chronological and ecological scope, and Russell Coope’s contribution A recent enumeration of records for the various data areas of BugsCEP is available elsewhere (http://www.bugscep.com; Buckland et al., 2014) and will not be repeated in full here. Similarly, a

Fig. 1. Simple classification of publications citing or using (either acknowledged or consulted) the Bugs database 1997e2012. (Note that this list includes 28 publications co-authored by a member of the Bugs development team).

bibliography of Russell Coope’s publications is available at http:// www.bugscep.com/downloads/coope_bibliography_2013.pdf. Buckland (2007) presented an overview of the geographical and chronological extent of the Bugs database, which is updated here to reflect the expanded database, with an indication of the extent to which Coope’s work is directly represented in the database. Note that the figures cited here to an extent reflect the way in which data has been entered into the database, and that there will inevitably be gaps, in particular with respect to the dating evidence which is often either difficult to source or insufficiently detailed to relate to a particular sample. A number of unpublished sites and reports attached to obscure archaeological monographs are missing, and the compilers would be happy to make these datasets publicly available through the database. 3.1. Geographical extent At the time of writing, the database contains raw data and metadata for 1135 sites, and Russell Coope directly contributed to the analysis of over 154 (14%) of them (Fig. 2); several North American sites and one on South Georgia (Coope, 1963) are not included. This represents some 98790 identifications published in 231 papers, some of which are explored further below. Connecting BugsCEP to GIS software or exporting query results for GIS use, increases the descriptive and analytical power of the database considerably, especially in terms of quantifying the spatial extent of particular aspects of the fossil record. Advanced queries may be created in the Microsoft Access backend for linking data on modern distribution and rarity with the geographical data in the fossil record (Fig. 3). By examining the dating and spatial evidence for particular species, it is possible to trace the movement of pests and synanthropic species (e.g. Bain and King, 2011). The reference data in BugsCEP may also be used to model potential past and future species movements with respect to climate change (e.g. Vickers and Buckland, in press). In addition to studies of biogeography, GIS techniques may be also applied to biographical questions. For example, the geographical mid-point of all sites on which Coope worked is latitude 52.7000, longitude
0.1224, just north of Whipchicken Road on the edge of Crowland in Cambridgeshire, UK (https://maps.google.com/maps? q¼52.7000386079137,þ-0.122489305971223&hl¼en&sll¼52.7000 39,-0.122489&sspn¼0.019921,0.028281&doflg¼ptk&t¼m&z¼15). The mid-point for his British Isles sites is somewhat more telling, just west of Kegworth, some 60 km northeast of Birmingham (https:// maps.google.com/maps?q¼52.8342843896639,þ-1.295575460243 7andhl¼enandll¼52.955257,-1.175537andspn¼1.267479,1.809998 andsll¼52.700938,-0.100937andsspn¼0.318725,0.452499andt¼ma ndz¼9). The spatial distribution of both modern and fossil insect collection sites may tell us more about where entomologists live and work than it does about the modern or fossil record, and the pitfalls of using these incomplete and heavily biased data should always be considered when drawing conclusions on the past and present distribution of species. It is only through the application of sound ecological knowledge that meaningful inferences can be drawn from such large amounts of data, however tempting large scale quantifications, such as some of those presented in this paper, may be. BugsCEP primarily contains data on the European fossil insect record, although it includes an increasingly large number of extraEuropean sites (ca. 100; see Buckland et al. 2014 for more details). The dominance of the British Isles data is clearly visible (Fig. 2; 58% of sites), and would undoubtedly skew any large scale analysis in the direction of the habitats favoured by the past and present fauna of these islands (see below). This bias would also inevitably lead to problems in geostatistical analyses, such as surface interpolation of environmental variables using kriging.

Please cite this article in press as: Buckland, P.I., The Bugs Coleopteran Ecology Package (BugsCEP) database: 1000 sites and half a million fossils later, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2014.01.030

P.I. Buckland / Quaternary International xxx (2014) 1e11

3

Fig. 2. Geographical extent of Coope sites (þ) against a background of other European sites in BugsCEP (dots).

Patterns at the small scale would inevitably be over represented as a result of the tight packing of sites in the UK, resulting in data artefacts such as anomalous climate reconstructions, in the areas where sites are sparse, such as northern Scandinavia or even the North Sea. The centre of gravity for all sites in the database is 40 km off the eastern coast of East Anglia, a point which shifts to the north of Germany with removal of the British sites. Fossil insect databases are under development for at least North American (Grimm et al., 2013), Russian (Sher et al., 2005) and Japanese (e.g. Shiyake, 2013) faunas, and the integration of international datasets should be considered a priority goal. Not only for increasing the data sharing and re-use capacity of insect analyses, but also in providing a broader ecological and climatic base for analyses. 3.2. Chronological extent

Fig. 3. Map showing the relative proportion of five habitats as represented at fossil sites by beetle species with Holocene records but now extinct in the UK. The underlying data were extracted from BugsCEP using a set of queries connecting the fossil record to a list of extinct species compiled from red data book data and a manual assessment of distribution data in the database. The map was then produced in ESRI’s ArcMap, and the list of species published as a section in the current Checklist of British Beetles (Duff 2012). Note the large proportion of sites where indicating the presence of species requiring “Wood and tree” habitats, which have now become extinct, possibly an indication of the loss of mid-Holocene woodland habitats, although the details of individual sites and species need further analysis.

By necessity of the disparate nature of the data sources behind the fossil insect record, dating evidence is varied in its form, accuracy and reliability. To cater for this, and to provide the capacity for storing data as recorded in a variety of sources, BugsCEP employs three categories for the storage of dates: 1) radiometric, including anything relying on a decay curve (Fig. 4; e.g. 14C, AAR, Useries); 2) period dates, where only broad determination of a sample to cultural, vegetation or geological period has been possible (Fig. 5; e.g. oxygen isotope stages, Blytt and Sernander vegetation zones, archaeological periods); and 3) calendar, including anything dated to a specific year or range of calendar years (Fig. 6; e.g. coin or other artefact-based dates, historical events, dendrochronology). Whilst every effort has been made to ensure the reliability of these dates, there are inevitably a large number of omissions and generalisations, much of which derives from the inconsistent reporting of dates in publications. As a consequence, the broad period dates have been used to patch holes and make the datasets more generally useful for looking at the wider chronological aspects of species distributions.

Please cite this article in press as: Buckland, P.I., The Bugs Coleopteran Ecology Package (BugsCEP) database: 1000 sites and half a million fossils later, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2014.01.030

4

P.I. Buckland / Quaternary International xxx (2014) 1e11

Fig. 4. Histograms showing the frequency of dates in BugsCEP. A) 14C over the last 15 000 years and B) the most recent 95% of radiometric dates. Coope sites are shown as a subset with striped subset; dates with only calibrated ages have been omitted for clarity.

The radiometric dates, and especially the 14C dates, held in BugsCEP, reflect the propensity of the palaeoentomological community to work on Holocene sites (Fig. 4). Clusters of dates are found in the Early and Late Holocene, and apart from the obvious explanation in the availability of material this pattern may partly be a result of the tendency to date the upper and lower parts of cores or peat sequences. BugsCEP does not yet include interpolated dates or calculate chronologies and relies on the use of broad period classifications as a workaround for providing contiguous overviews. Coming versions of the software will include simple calibration of 14 C dates to enable rapid searching and comparison across sites, as well as more complete chronologies. Coope’s interest in the Late Glacial and earlier periods is clearly illustrated by the relative frequency of dates from his sites in the database. In fact, 77% of all radiometric dates older than 10 000 BP come from Coope sites. The importance of the Late Glacial and Holocene for the Quaternary insect record is similarly illustrated by the frequency of samples classed as such using the broad period categories of BugsCEP (Fig. 5). These categories are used to improve searching facilities when comparing the fauna of different sites and samples, a simple surrogate for implementing chronologies for almost 1000 sites, some of which have no accepted chronology. The period based system is also necessary for many archaeological sites where absolute dating evidence is either poor, lacking, or established by broad relative typologies. This system is, however, only partially implemented in that it is not currently possible to easily search for multiple periods or to automatically include sub-periods in a search for an encompassing period (e.g. Loch Lomond within Late Glacial).

In SEAD, the use of a period dictionary will alleviate this problem by chronologically defining period boundaries and enabling equivalents to be automatically compared in searches. Where relevant, period definitions will also be spatially referenced in order to cater for the lag in the geographical spread of archaeological cultural periods. A large number of archaeological sites have on the other hand the advantage of accurately datable material for providing calendar dates, such as coins, pottery, or dates from the historical record, for example, the faunas preserved by the eruption of Vesuvius in AD 79 (dal Monte, 1956) or those from forts on the Antonine Wall in Scotland, occupied ca. AD 139e160 (Smith, 2004). These methods, when combined with the complexity of excavations and the need to date contexts by association or stratigraphic relationship, often lead to samples representing a range of between two established constraints (or termini; Fig. 6). To extract these data for more than the specifically dated contexts often requires delving deep into site archives, and details are frequently poorly reported in journal publications. They are therefore of variable reliability for high precision studies and users are required to consult the primary literature before drawing too definite conclusions. Frequency of dates is not a good measure of past cultural activity, and although it has occasionally been used to imply past population or site densities, the statistic is largely dependent on other factors, including archaeological interest for the period or culture, site preservation, and variations in excavation, dating strategy and sampling methodology. Two peaks are particularly prominent in Fig. 6, resulting from archaeoentomological interest in the medieval period (ca. 1000e400 BP) and Roman Britain (ca. 2000e1500 BP). These are of

Please cite this article in press as: Buckland, P.I., The Bugs Coleopteran Ecology Package (BugsCEP) database: 1000 sites and half a million fossils later, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2014.01.030

P.I. Buckland / Quaternary International xxx (2014) 1e11

5

Fig. 5. Bar chart showing number of samples classed as belonging to each chronological, cultural or historical period. Note that these periods need revision and more thoroughly defining chronologically and geographically in order to provide a reliable source for biogeography studies. Samples from Coope sites are shown as a striped subset.

course not only popular periods for research, but benefit from abundant datable material and to an extent corroboration historical records. 4. Analysis tools Whilst a relational database allowing comparisons between the Quaternary fossil record and modern habitat and distributional

data has considerable value, it is further enhanced by the addition of tools for data analysis. Simple data features such as synonymy, species associations and some keys, part of an uncompleted project by the late Peter Skidmore, who incidentally also provided the illustration for the software’s front page, with additional comments on identification, and size range of species extend the work into a desk-based system to assist in basic laboratory work. The whole is supported by an extensive bibliography of over 5000 references.

Fig. 6. Histogram showing frequency of calendar date termini (from, to) by age BP (present ¼ 1950 for comparative purposes). The inset shows the same data but plotted as ranges between dating termini.

Please cite this article in press as: Buckland, P.I., The Bugs Coleopteran Ecology Package (BugsCEP) database: 1000 sites and half a million fossils later, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2014.01.030

6

P.I. Buckland / Quaternary International xxx (2014) 1e11

Whilst Coope had a seemingly unerring ability to retrodict contemporary climate from a fossil insect assemblage, it was the need to place numerical parameters around such apparently accurate inferences that lead to the development of MCR, mutual climatic range. Initially devised by Atkinson, Briffa and Coope (1986) and expanded by Joachim and Perry (Atkinson et al., 1986, 1987), the module has now been re-written and incorporated within BugsCEP (Buckland, 2007). 4.1. Climate reconstruction Essentially, MCR (Mutual Climatic Range) uses geographic distribution to define the climate space occupied by species in terms of two defining parameters, mean summer temperatures, and annual temperature range, the latter effectively a measure of continentality. These are quantified from modern and historical collection data into species envelopes in terms of TMax, mean temperature of the warmest month and TRange, the difference between TMax and TMin, mean temperature of the coldest month. TRange may also be considered an index of continentality. Individual species’ climate envelopes are then overlain to define an area of mutual climate range, although not necessarily co-occurrence, for the species found in a sample. Currently data only include the original envelopes for carabids, staphylinids, scarabaeids and a few other groups painstakingly put together and digitised by Joachim and Perry in the days when Fortran was the only language capable of dealing with the numerics involved, and there is a desperate need to update the basic information, utilising a more advanced GIS and primary specimen data. As national databases of modern distribution have become more readily available (e.g. Luff, 1998; Cox, 2007 for UK examples; Artdatabanken, 2013 for Sweden), this task has become more easily accomplished both in terms of refining envelopes and expanding to other groups not only of Coleoptera but also other frequent insect fossils such as the Formicidae, ants. The development of international, point source database systems for collection and museum data, such as the Global Biodiversity Information Facility (GBIF, 2013), although as yet unreliable for the task (see e.g. Buckland and Eriksson, 2014), should also make the future construction of envelopes easier. Despite the limitations imposed by reliance upon often poor primary data, MCR has been routinely applied to Pleistocene assemblages for over twenty years, and its basic Lyellian philosophy has attracted Quaternary scientists working with other groups, including plant macrofossils (Pross et al., 2000; Thompson et al., 2012), terrestrial mollusca (Moine et al., 2002) and ostracoda (Horne, 2007). In addition, it has been applied to assemblages in other geographical regions, including Beringia (Alfimov and Berman, 2009) and Japan (Shiyake, 2013). The application of the method, however, has often had problems which are difficult to overcome, relating more to context taphonomy and minimum species diversity per sample than distributional data. The stochastic nature of sampling means that the same context may provide either a broad date range from a single relatively eurythermal species or a narrow one from a stenotherm. These problems are, however, common to all quantitative reconstruction methods, and are reduced by data and method transparency. For this reason, BugsCEP displays the species and envelopes being utilised for the MCR reconstruction, and does not apply the now questioned calibration technique used to derive mean temperatures in earlier studies (see Buckland, 2007; Elias, 2010). Standardisation of sample size across a varying sequence of sediments may appear statistically sound, but may be of little real value (see below). In the case of open exposures, as in Coope and

Brophy’s (1972) Glanlynnau work, long before the days of MCR, the answer was to go back and get more material, thereby creating a more useful reconstruction both in terms of climate and environment (Fig. 7). The resultant summer temperature curve, drawn by Coope as a guestimate mid-line is remarkably similar to that obtained by MCR and both emphasise the step-like nature of the transition from cool (July average w11  C) to warm (w17  C) summers, something which involved him and co-workers in seemingly endless disagreements, particularly with palynologists, until similar evidence for rapid transitions emerged from the Greenland ice cores some twenty years later (cf. Alley et al., 1993). BugsCEP allows the generation of similar MCR diagrams in a matter of seconds, and Fig. 8, utilising Osborne’s (1980) West Bromwich sequence, shows the similar abrupt transition into the Holocene in its raw, unscaled, output form with jackknife limits added from the advanced output. Whilst MCR is ideal for the broad brush treatment of glacial, interglacial, stadial interstadial climate, it currently lacks the detail to apply to examine later Holocene climate change, where human impact becomes increasingly the dominant factor, and more reliable envelopes are required for a broader range of species. Robinson (2013) has recently argued for a short period of significantly warmer summers during the Bronze Age on the basis of dung faunas from sites in southern England. MCR on sites such as Stanwick in Northamptonshire (Robinson, 2013) and Wilsford in Wiltshire (Osborne, 1989) would appear to support this, although there are still too few dated sites to differentiate effectively between human impact and climate change. In the case of the dung fauna, the ploughing up of old grassland and more recently the use of ‘biocides’ on domestic stock has significantly altered the fauna to the extent that reliable habitat data for some species relies heavily on older records. Climate reconstruction methods can often be turned their heads to model potential biogeographic impacts of future, or past climate scenarios. BugsCEP provides simple tools for listing the species which would be able to survive in any given temperature range, or which species could theoretically co-exist in the same climate given unlimited migration possibilities. Again, the limits here are the quality and extent of the calibration data, with only 436 species in the MCR calibration dataset. It is however, possible to do a reasonable amount of modelling on this basis, albeit with a degree of back-end hacking, and the system lends itself well to, for example, evaluating existing theories on island biogeography and species survival over varying climate periods (e.g. Vickers and Buckland, in press). 4.2. Habitat data, reconstruction, comparing and standardising across samples A notorious problem in ecology and palaeoecology alike is the establishment of a frame of reference for comparing samples of different size (volume, weight, spatial and chronological extent), collected using different methods (bulk samples, stratigraphic sampling, cores, pitfall traps), from different depositional environments (lakes, bogs, archaeological sites) and representing different environments with differing population structures and sedimentation rates. These differences are accompanied by different of sampling and taphonomic problems which must be accounted for, or at least considered, when interpreting the environmental implications of the fossil insect fauna. Raw minimum numbers of individuals (MNI) most often tells us more about the nature of the samples themselves than the environments from which their fossils originated. These problems have been discussed with respect to palaeoentomology and

Please cite this article in press as: Buckland, P.I., The Bugs Coleopteran Ecology Package (BugsCEP) database: 1000 sites and half a million fossils later, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2014.01.030

P.I. Buckland / Quaternary International xxx (2014) 1e11

Please cite this article in press as: Buckland, P.I., The Bugs Coleopteran Ecology Package (BugsCEP) database: 1000 sites and half a million fossils later, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2014.01.030

Fig. 7. Climate and habitat reconstructions at A) Glanllynnau Coope and Brophy (1972) and B) Abisko (modern dataset, current author). The pre-MCR summer temperature curve of Coope and Brophy (1972) is overlaid on the equivalent ranges produced by BugsCEP. (Note that this figure has been scaled by depth and visually improved, using Golden Software’s Grapher 9, with respect to the Excel raw output provided by BugsCEP (see Figure G for an example). Excel output is a simple solution for connectivity, but far from ideal in terms of visualisation and manipulation ease). The habitat proportions are based on finds identified to species level, presence data only and calculated according to the BugStats method described by Buckland (2007), standardised by total number of habitat indications per sample (see Buckland, 2007 for full method). MNI ¼ Minimum Number of Individuals.

7

8

P.I. Buckland / Quaternary International xxx (2014) 1e11

Fig. 8. BugsMCR diagrams for the West Bromwich sequence (Osborne, 1980), dated to between 12 195  160 (Sample A) and 9080  455 (Sample M) 14C BP. The black floating boxes with whiskers show the area of maximum overlap (box) and extreme extent resulting from removing any one species from the MCR calculations (whiskers). The rapid shift in summer temperatures is seen between samples E and F. The pale floating boxes indicate MCR ranges derived from less than three species and without jackknife calculations. The number of species used in the MCR calculations is shown by the wide bars on the TMax graph.

archaeoentomology by several authors (e.g. Carrot and Kenward, 2001; Buckland, 2007) and a number of approaches proposed to deal with them. BugsCEP provides tools for aiding the comparison of samples, both in terms of species numbers and the environmental implications of the species found at any site. It does not, however, provide the facility to standardise by sample weight or volume, but instead focusses on the numbers of taxa, individuals and habitat statistics (see Buckland, 2007 for full method description). Fig. 7 shows habitats standardised by the total count of habitat indications for each sample. This reduces the effects of varying numbers of species and/or individuals between samples, which in the case of highly variable sites, like Glanllynnau, would overpower any environmental signals in the data, and tend to mirror the observation data (observe the form of the MNI diagram in Fig. 7). In this example, only numbers of species are used, essentially illustrating the environmental implications of the species present in each sample. The software can also be set to include abundance data in the calculations, in order to examine the importance of varying numbers of individuals between samples and habitats, or compare samples containing comparable numbers of individuals. Higher level taxa (e.g. Aphodius sp.) may also be included to generalise the reconstruction, a useful facility for sites with poor preservation and where only a broad environmental description is required. The system can be used to look at fossil and modern assemblages alike, as suggested by the inclusion of the Abisko reconstruction at the bottom of Fig. 7, and thus provides a mechanism for the quantitative comparison of these fauna. Interestingly, the modern Abisko fauna gives a climate reconstruction similar to the colder parts of the Glanllynnau sequence, as expected, but Abisko’s dwarf shrub/wetland environment appears more to be more similar to the warmer parts (the data for the “Abisko modern” reconstruction are available in BugsCEP. The environmental reconstruction function is provided as an investigative tool, not a system for producing answers, in palaeoenvironmental reconstruction. It allows users to reduce the

sensitivity of their reconstructions to variations in species numbers caused by variations in sample size, preservation or population richness. The system of database coupled habitat codes (ecocodes) and standardisation methods (BugStats) is also designed to allow samples to be compared between sites irrespective of author, and thus allows for more objective inter-site overviews than are easily achievable when comparing texts. It should not be forgotten, however, that variation in the number of fossils per unit sample volume is a product of complex environmental, biological and taphonomic processes which we should attempt to understand in terms of habitat, sedimentology and sampling when reconstructing the past. 4.3. Exploring data and the variable representation of environments Databases provide unique possibilities for exploring and quantifying variability in the empirical evidence behind palaeoenvironmental reconstructions. The under-representation of insect species in the fossil record is perhaps illustrated by the fact that of the 10,546 taxa for which the database holds reference data, only 4418 have a published fossil record (Table 1; assuming at least a 90% coverage of the published Quaternary fossil record for Europe). Such summary numbers are, however, potentially deceptive as the selection of reference taxa is often a function of the data entry strategy for the database, some hidden implications of which are discussed below. Although a little used facility, the database may also be used to quantify information from a bibliographic perspective, as Table 1 illustrates for the data contribution of Coope. This line of enquiry could potentially be combined with the above overview of dating and geographical evidence (Figures C, D and E), as well as citation records, for compiling research histories, looking at trends in the interests and foci of individuals over their careers and the re-use and persistence of data, something which is particularly evident in the habitat data for the rarer species, where the only captures in a country may have been made over a century ago and the primary data are reiterated by later authors.

Please cite this article in press as: Buckland, P.I., The Bugs Coleopteran Ecology Package (BugsCEP) database: 1000 sites and half a million fossils later, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2014.01.030

P.I. Buckland / Quaternary International xxx (2014) 1e11 Table 1 Statistics from the database: simple representivity of the fossil record and a simple quantification of Russell Coope’s direct contribution to the data. To put it in perspective, there are ca 200 individuals credited with having identified fossil material in the database (although this list is incomplete and somewhat unreliable in its current form). Fossil

Modern

%

Number of taxa in database

4418

10546

41.9

Coope

Total in BugsCEP

%

Unique sites with Coope IDs Fossil publications co-authored by Coope Fossil publications with Coope IDs Unique taxa ID’d by Coope .of which to species level Total identifications ...of which to species level Species level ID’s as % of sum of Total ID’s

154 132 231 1922 1331 98790 61634 62.4

1124 1193 1193 10546 8367 602480 356071 59.1

13.7 11.1 19.4 18.2 15.9 16.4 17.3

Although interesting in terms of research history, the above numbers tell us little directly about the representivity of species groups, and by implication, habitats in the database. When using databases there is often an assumption that the database contains an objective, neutral mass of data which may be safely handled without looking at the extent and quality of its coverage. (This is perhaps why a number of organisations seem to be under the impression that we need now only build portals to link databases in order to solve most current research questions). BugsCEP is of course no exception, and the data contained within are highly biased as a result of the research history of the database authors, the data contributors, their (limited) linguistic skills, and the availability of reference data for different species. Exactly how these biases impact on interpretations depends to a large extent on how data are extracted, manipulated and analysed. Fig. 9 further illustrates what is evident in the dating figures above but in terms of broad habitat classifications (see Buckland, 2007, p102 for a detailed description of the habitat classes and the rationale behind them). This essentially simple diagram contains an enormous amount of hidden information, a full explanation of which is outside the scope of this paper. Although superficially illustrative of faunal diversity, the diagram is fundamentally misleading, being highly dependent on the primary sources chosen for reference data,

9

and while these include 3043 publications, there are a number of larger compendia, such as Koch (1989e92), which undoubtedly skew the distribution towards their foci. The large number of “Wood and trees” taxa may be a representation of this particular source, which is based mainly on central European studies, although again this may also be influenced by the English Nature funded project on the nature of pre-forest clearance woodland (Hodder et al., 2005). Generally speaking, the observation and abundance figures (Fig. 9:B and C) mirror the figures for numbers of reference taxa per habitat, as would be expected when reference data are entered as needed to allow for the description of the fossil fauna. There are, however, a few interesting exceptions: The proportionally larger number of finds of species classed as preferring Wetlands/marshes could very well have a simple explanation: many samples are taken from peat successions and these habitats more often than not are dominated by a site specific fauna (e.g. Whitehouse, 2004). Species indicating Pasture/Dung, Dung/foul habitats and General synanthropic species are also overrepresented in the fossil finds, and this could be an indication of the archaeological aspect of the database (see Buckland et al., 2014), where the potentially wetter, anaerobic environments of cess pits, storage pits, wells, and house floors provide the best preservation. On a biographic note, it may be interesting to observe that Coope holds the records for the largest numbers of individuals of any species found in any sample representing Standing water, Running water, Meadowland and Heathland and moorland. This is however, no simple coincidence, but a reflection of his extensive work on large samples from Lateglacial deposits. Fig. 9:D provides in itself some useful information on the taphonomy and biodiversity of particular environments with respect to superabundant taxa. Further analysis would be required to find out whether these numbers reflect sample sizes or high relative abundances. It should also be pointed out that the average number of individuals per sample varies between 2.1 and 14.4, and that the extremes are anomalies which may need to be removed as outliers prior to any quantitative biodiversity analysis. The figure thus serves to illustrate another pitfall of uncritically using database sources. Some of these anomalies are a reflection of how the data have been aggregated into samples, either in publications or in BugsCEP,

Fig. 9. The absolute representation of broad habitat categories in BugsCEP. From left to right these are expressed, per category, as A) number of taxa in the database and number of these known fossil, followed by this relationship as a percentage; B) total number of observations, where one observation ¼ one recording of a taxon in a sample; C) total number of individuals recorded (MNI) and D) the maximum number of individuals of any single taxon recorded in a sample. Coope’s contribution is illustrated by the striped subset of diagrams BeD. Note the different scales and that a taxon may be found in >1 category.

Please cite this article in press as: Buckland, P.I., The Bugs Coleopteran Ecology Package (BugsCEP) database: 1000 sites and half a million fossils later, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2014.01.030

10

P.I. Buckland / Quaternary International xxx (2014) 1e11

pending full quantification or in absence of a full list of taxa per sample. The Lateglacial assemblage from Lobsigensee (Elias and Wilkinson, 1983), for example, produced almost 4000 unresolvable specimens of the aquatic genus Ochthebius. Examination of the sample metadata shows, however, that the data from this site are number of individuals per faunal unit, and not per physical sample. This number is topped only by the 8351 pupae of the (non-Bugs EcoCode classified) fly Trachyopella coprina (Duda) identified by the late Peter Skidmore from an Anglo-Scandinavian ditch at Tuquoy in Scotland (Skidmore, 1996). Aggregated data from Wabmer’s (1995) survey of modern pasture habitats also provides large peaks for the several dung related categories. In these cases it is perhaps the presence of particular species and the relative frequency of taxa within the site that are of most use to the palaeoentomologist, although the database needs many more quantified modern assemblages entering to allow effective comparison with fossil data. Robinson’s (2013) recent work with the scarabaeid dung fauna, whilst not utilising BugsCEP, points in the right direction. The BugsCEP software provides a variety of tools for searching almost any aspect of the database, including the ecology, rarity, distribution and temperature range components of the modern reference data. It also allows the user to explore the details of any BugStats based reconstruction, in both spreadsheet and report forms, in the spirit of total research transparency. Such facilities not only provide for powerful exploratory data analysis, but also increase the range of potential users beyond those interested in insects and at least into the realms of climate science, biodiversity and conservation studies. The ability to easily connect any of these results with data from the fossil record is an extremely powerful tool for helping to understand the origins of the modern landscape from any number of angles. 5. Current developments and multiproxy analyses In terms of software, although not data entry, the BugsCEP system is currently in limbo awaiting transfer to the Strategic Environmental Archaeology Database (SEAD; http://www.sead.se; e.g. Buckland, 2014). The data are being ported over at the time of writing and soon will be available online. Having been written for now obsolete versions of both Microsoft Office and Windows, BugsCEP is becoming increasingly difficult to run on new computers, and the authors have little time available for the continual maintenance required to fix problems caused by endless Windows updates. The system is a complex patchwork of program modules totalling approximately 70 000 lines of code, and it will inevitably take some time before the full functionality is realised on a web based platform. For this reason BugsCEP will be supported for as long as possible in parallel with SEAD. Moving Bugs onto the web provides scope for a number of substantial improvements in terms of data entry, accessibility and linking with other datasets (see Buckland, 2014; Buckland and Eriksson, 2014). Projects are underway in Sweden as part of SEAD to connect the insect data, and in particular the results of environmental reconstructions, to spatially referenced cultural heritage data. The long term plan with this is to provide a system which will be able to provide online graphical landscape reconstructions, based on real data, for areas around national monuments. In addition to insect data, SEAD will provide data on plant macrofossils, pollen, geochemistry, dendrochronology and ceramics to allow for a more comprehensive reconstruction of past cultural and natural environments. There are any number of hurdles to cross on the way to fulfilling this vision, not least the collection and digitisation of more empirical data e the keystone of all science. The Bugs database is also scheduled for mirroring in the international Neotoma database (http://www.neotomadb.org; Grimm et al.,

2013), a move which will greatly enhance the capacity for multiproxy and international syntheses. 6. Conclusions BugsCEP has come a long way since its humble, yet groundbreaking origins, and is scheduled for even greater things. The inclusion of tools for faunal and environmental analysis has greatly improved its applicability to areas of science far beyond the inherently multidisciplinary fields of Quaternary entomology and archaeoentomology. Although powerful both in terms of the large mass of aggregated data, which allows for large scale spatial and chronological analyses, and in terms of integrating modern and fossil data, the database and software should only be used in the context of a detailed knowledge of insect ecology if reliable conclusions are to be drawn through its use. Acknowledgements Like all long term projects, BugsCEP builds on its humble precursors, devised initially by Jon Sadler and Mike Rains. Paul Buckland has been responsible for suggesting numerous features, without which the system would be little more than an archive, as well as entering the bulk of the data. Data entry is an underappreciated and often massively under-budgeted component of any database project, and it is only as a result of Paul’s decades of tireless work that the database has been sustained. Corrections and additions to the contents of database should be directed to Paul. As demonstrated in this article, Russell Coope is responsible for the creation of the largest subset of these data. Through a more complex process, involving genetics and undoubtedly cats, he is also indirectly responsible for turning a young programmer into a young palaeoentomologist and thus the current form of the software. Finally, a database is nothing without its data contributors, who are individually acknowledged through their records in the database’s bibliography. References Alfimov, A.V., Berman, D.I., 2009. Possible errors of the Mutual Climatic Range (MCR) method in reconstructing the Pleistocene climate of Beringia. Entomological Review 89 (4), 487e499. Alley, R.B., Meese, D.A., Shuman, C.A., Gow, A.J., Taylor, K.C., Grootes, P.M., White, J.W.C., Ram, M., Waddington, E.D., Mayewski, P., Zielinski, G.A., 1993. Abrupt increase in the Greenland snow accumulation at the end of the Younger Dryas event. Nature 362, 527e529. Artdatabanken, 2013. The Swedish Species Information Centre. URL: http:// www.slu.se/en/collaborative-centres-and-projects/artdatabanken/species/ (last accessed 08.07.13.). Atkinson, T.C., Briffa, K.R., Coope, G.R., Joachim, M.J., Perry, D.W., 1986. Climatic calibration of coleopteran data. In: Berglund, B.E. (Ed.), Handbook of Holocene Palaeoecology and Palaeohydrology. John Wiley and Sons Ltd, London, pp. 851e 858. Atkinson, T.C., Briffa, K.R., Coope, G.R., 1987. Seasonal temperatures in Britain during the past 22,000 years reconstructed using beetle remains. Nature 325, 587e 592. Bain, A., King, G., 2011. Asylum for Wayward immigrants: historic ports and colonial settlements in Northeast North America. Journal of the North Atlantic 1, 109e 124. Buckland, P.I., 2007. The Development and Implementation of Software for Palaeoenvironmental and Palaeoclimatological Research: the Bugs Coleopteran Ecology Package (BugsCEP) (PhD thesis). In: Archaeology and Environment, vol. 23. Environmental Archaeology Lab. Department of Archaeology and Sámi Studies. University of Umeå, Sweden. URL: http://urn.kb.se/resolve?urn¼urn: nbn:se:umu:diva-1105. Buckland, P.I., 2014. SEAD - The Strategic Environmental Archaeology Database. Inter-linking multiproxy environmental data with archaeological investigations and ecology. In: Earl, G., Sly, T., Chrysanthi, A., Murrieta-Flores, P., Papadopoulos, C., Romanowska, I., Wheatley, D. (Eds.), CAA2012, Proceedings of the 40th Annual Conference of Computer Applications and Quantitative Methods in Archaeology (CAA). Southampton, England. Amsterdam. Buckland, P.I., Buckland, P.C., 2006. BugsCEP Coleopteran Ecology Package. In: IGBP PAGES/World Data Center for Paleoclimatology Data Contribution Series #

Please cite this article in press as: Buckland, P.I., The Bugs Coleopteran Ecology Package (BugsCEP) database: 1000 sites and half a million fossils later, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2014.01.030

P.I. Buckland / Quaternary International xxx (2014) 1e11 2006-116. NOAA/NCDC Paleoclimatology Program, Boulder CO, USA. URL: http://www.ncdc.noaa.gov/paleo/insect.html http://www.bugscep.com. Buckland, P.I., Buckland, P.C., Olsson, F., 2014. Paleoentomology: insects and other arthropods in environmental archaeology. In: Smith, C., Lanteri, C., Reid, J., Smith, J., Krauss, T.M. (Eds.), The Encyclopedia of Global Archaeology. Springer. Buckland, P.I., Eriksson, E.J., 2014. Strategic environmental archaeology database (SEAD). In: Smith, C., Lanteri, C., Reid, J., Smith, J., Krauss, T.M. (Eds.), The Encyclopedia of Global Archaeology. Springer. Buckland, P.I., Eriksson, E.J., Linderholm, J., Viklund, K., Engelmark, R., Palm, F., Svensson, P., Buckland, P.C., Panagiotakopulu, E., Olofsson, J., 2010. Integrating human dimensions of Arctic palaeoenvironmental Science: SEAD e the strategic environmental archaeology database. Journal of Archaeological Science 38, 345e351. Carrot, J., Kenward, H., 2001. Species, associations among insect remains from urban archaeological deposits and their significance in reconstructing the past human environment. Journal of Archaeological Science 28, 887e905. Coope, G.R., 1963. The occurrence of the beetle Hydromedion sparsutum (Müll.) in a peat profile from Jason Island, South Georgia. British Antarctic Survey Bulletin 1, 25e26. Coope, G.R., Brophy, J.A., 1972. Late Glacial environmental changes indicated by a coleopteran succession from North Wales. Boreas 1, 97e142. Cox, M.L., 2007. Atlas of the Seed and Leaf Beetles of Britain and Ireland (Coleoptera: Bruchidae, Chrysomelidae, Megalopodidae and Orsodacnidae. Pisces, Newbury. dal Monte, G., 1956. La presenza di insetti dei granai in frumento trovato neg1i scavi di Ercolano. Redia 41, 23e28. Duff, A. (Ed.), 2012. Checklist of beetles of the British Isles. Elias, S.A., 2010. Advances in Quaternary Entomology. In: Developments in Quaternary Sciences, vol. 12. Elsevier. Elias, S.A., Wilkinson, B., 1983. Lateglacial insect fossil assemblages from Lobsigensee (Swiss Plateau). Studies in the Late Quaternary of Lobsigensee 3. Revue de Paléobiologie 2, 189e204. GBIF, 2013. The Global Biodiversity Information Facility. URL: http://www.gbif.org/ (last accessed 08.07.13.). Grimm, E.C., Bradshaw, R.H.W., Brewer, S., Flantua, S., Giesecke, T., Lézine, A.-M., Takahara, H., Williams, J.W., 2013. Databases and their application. In: Elias, S.A. (Ed.), The Encyclopedia of Quaternary Science, vol. 3. Elsevier, Amsterdam, pp. 831e838. Hodder, K.H., Bullock, J.M., Buckland, P.C., Kirby, K.J., 2005. Large Herbivores in the Wildwood and Modern Naturalistic Grazing Systems. English Nature, Peterborough. Horne, D.J., 2007. A Mutual Temperature Range method for Quaternary palaeoclimatic analysis using European nonmarine Ostracoda. Quaternary Science Review 26 (9e10), 1398e1415.

11

Koch, K., 1989-92. Ökologie IeIII. Die Käfer Mitteleuropas. Goecke and Evers, Krefeld. Luff, M.L., 1998. Provisional Atlas of the Ground Beetles (Coleoptera, Carabidae) of Britain. Centre for Ecology and Hydrology, Monks Wood. Moine, O., Rousseau, D.-D., Jolly, D., Vianey-Liaud, M., 2002. Paleoclimate reconstruction using Mutual Climatic Range on terrestrial mollusks. Quaternary Research 57, 162e172. Osborne, P.J., 1980. The Late Devensian Flandrian transition depicted by serial insect faunas from West Bromwich, England. Boreas 9, 139e147. Osborne, P.J., 1989. Insects. In: Ashbee, P., Bell, M., Proudfoot, E. (Eds.), Wilsford Shaft Excavations 1960e62. English Heritage, London, pp. 96e99. Pross, J., Klotz, S., Mosbrugger, V., 2000. Reconstructing palaeotempertures for the Early and Middle Pleistocene using the mutual climatic range method based on plant fossils. Quaternary Science Reviews 19, 1785e1799. Robinson, M., 2013. The relative abundance of Onthophagus species in British assemblages of dung beetles as evidence of climate change. Environmental Archaeology 18, 132e142. Sadler, J.P., Buckland, P.C., Rains, M., 1992. BUGS: an entomological database. Antenna 16, 158e166. Sher, A.V., Kuzmina, S.A., Kuznetsova, T.V., Sulerzhitsky, L.D., 2005. New insights into the Weichselian environment and climate of the East Siberian Arctic, derived from fossil insects, plants, and mammals. Quaternary Science Reviews 24 (5), 533e569. Shiyake, S., 2013. Applying the Mutual Climatic Range method to the beetle assemblages in Japan using accurate data of climate and distribution of modern species. Quaternary International. http://dx.doi.org/10.1016/j.quaint.2013.06.002. Skidmore, P., 1996. A Dipterological Perspective on the Holocene History of the North Atlantic Area (Unpublished PhD thesis). University of Sheffield. Smith, D.N., 2004. The insect remains from the well. In: Bishop, M.C. (Ed.), Inveresk Gate: Excavations in the Roman Civil Settlement at Inveresk, East Lothian, 1996e2000, STAR Monograph 7, pp. 81e88. Loanhead, Midlothian. Thompson, R.S., Anderson, K.H., Pelltier, R.T., Strickland, L.E., Bartlein, P.J., Shafer, S.L., 2012. Quantitative estimation of climatic parameters from vegetation data in North America by the mutual climatic range technique. Quaternary Science Reviews 51, 18e39. Vickers, K., Buckland, P.I. Predicting island beetle faunas by their climate ranges e the tabula rasa/refugia theory in the North Atlantic. Journal of Biogeography (in press). Waßmer, T., 1995. Mistkäfer (Scarabaeoidea et Hydrophilidae) als Bioindikatoren für die naturschützerische Bewertung von Weidebiotopen. Zeitschrift für Ökologie und Naturschutz 4, 135e142. Whitehouse, N., 2004. Mire ontogeny, environmental and climatic change inferred from fossil beetle successions from Hatfield Moors, eastern England. The Holocene 14, 79e93.

Please cite this article in press as: Buckland, P.I., The Bugs Coleopteran Ecology Package (BugsCEP) database: 1000 sites and half a million fossils later, Quaternary International (2014), http://dx.doi.org/10.1016/j.quaint.2014.01.030