BULLETIN DE L'INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE ENTOMOLOGfE, 79: 35-44, 2009 BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN ENTOMOLOGJE, 79: 35-44, 2009 Carabid beetles as ecological indicators in Belgian archaeology: the pioneering work of Konjev Desender By Anton ERVYNCK Summary interaction with the enviromnent. In fact, it became clear that humans not only cannot exist or smvive An overview is given of the development of the analysis of without exploiting other species (animal and plant archaeological carabid remains in Belgium, and of the crucial resources) but also that much (most, even everything role played by Konjev Desender. The bibliography of the sites following the viewpoint of 'ecological determinism') investigated over the last decennia is presented, and explanations are put forward for the rather biased nature of the dataset collected. within human behaviour was and is influenced by the An evaluation of the interpretive value of this research line indicates characteristics of the surrounding landscape, geology, that the integration of the archaeological data into the recent soil, flora and fauna. Although the analysis of animal biodiversity databases is a major step forward. and plant remains as part of archaeological research is already an old phenomenon (see the contributions Key words: Carabidae, Belgium, archaeology, ecological recon structions by botanists and zoologists, in the middle of the 19th century, to the excavations of Danish shellmiddens Resume and the Swiss lakeside dwelling sites), the idea that the human species was part of a local, regional or global Cet article rend compte du developpement de Ia recherche sur les ecosystem, only gained considerable attention during restes archeologiques de coleopteres carabiques en Belgique et du the second half of the 20th century (see TRIGGER, 2006, role essentiel qu'a joue Konjev Desender dans ce processus. Une for a history of archaeology). bibliographie des sites ou les carabes ont ete etudies est etablie et les raisons de Ia nature peu equilibree du set de donnees sont expliques. This development invoked a stimulus for the analysis Apres !'evaluation de Ia valem interpretative de ces recherches il est of non-cultural material derived from excavations. clair que !'integration des donnees archeologiques dans les bases de Gradually, more time and effort was invested into the donnees de bio-diversite recentes est un grand pas en avant. sampling and recovery of organic finds present in the soil and deposits that constitute the . 'archaeological Mots-clefs: coleopteres carabiques, Belgique, archeologie, analyses environnementales record'. This trend resulted in the growth of a specialist branch within archaeology, 'environmental archaeology', concentrating on the interaction between Introduction humans, plants and animals in the past, through the study of former landscapes, (buried) soils, botanical Archaeology is the science that reconstmcts, interprets and zoological remains (see EvANs & O'CoNNOR, 1999 and explains human behaviom in the past, through the for an introduction). The growing attention towards the analysis of excavated material remains. Originally, recovery of organic remains dming excavations led the conceptual framework was exclusively cultmal, to the implementation of sieving techniques allowing as if the development of the hwnan species was only adequate sampling of even the minutest biological a matter of ideas, rituals, beliefs, politics, other social remains. Within these finds collections, material from interactions, etc. However, with the rise of the 'New small animals was sometimes abundant and its presence Archaeology', in the '60s of the 20th century, the idea could often not be explained by hwnan conswnption became widely accepted that former human behaviour patterns, industrial or small scale use of animal products cannot be lmderstood without taking into account or other activities. Indeed, the extensive sieving - '' 36 A. ERVYNCK campaigns showed that, in the past, a wide variety of not yet published (unfortunately, considerable delays smaller animals had ended up, without any direct human in the publication of excavation reports are a true interference, in what are now archaeological contexts. characteristic of archaeology). Those from another site, Amongst the remains of these so-called 'intrusives', not a Roman fortification at Maldegem, did find their way only small rodents, insectivores, amphibians, reptiles or into publication (MERTENS et a!., 1986) (see Fig. 1 for terrestrial molluscs are found, but also many species of the location of all sites mentioned). insects and other arthropods. Taking into the account the development of the whole Originally sometimes described as 'background field of environmental archaeology in the country, with fauna' (without too much importance for the study of the first study of animal remains only being published former human behaviour), the intrusive animal remains in 1965 and sieving becoming common only since are now widely used within archaeology as ecological the 1980s, the late date of this innovation seems indicators. Through an evaluation of the ecological understandable (see www.onderzoeksbalans.be for a characteristics of the intrusive species found, i.e. their detailed description of the development of archaeology habitat preferences (as known from the ecological in Flanders). Even in the U.K., one of the pioneering studies of actual biotopes), inferences can be made countries for environmental archaeology, the first about the former environment around the archaeological archaeological insect studies were only published in the context investigated. The basic assumption taken when second half of the 20th century (BucKLAND, 1976). attempting such ecological reconstructions is that the Despite the enormous potential of archaeological 'principle of actuality' is valid (commonly used in entomology, the practicalities of archaeology (the palaeontology and stating that "the present is the key to constant lack of adequate funding, the need for the past"), implying that the ecological characteristics expertise in many different fields) have hampered a full of a species should not have changed significantly exploitation of this research line. In a discussion of the through time. Given the evolutionary characteristics of use of the remains of mites (Acarina) in archaeology, insects, and the relatively short time lapse between the ScHELvis (1993) once compared the relative abundance actual fauna and its archaeological predecessors (when of animal groups within the living world against the evaluated on an evolutionary or geological scale), this research themes followed by archaeozoologists (Fig. 2) assumption seems safe to accept (CoorE, 1978). and concluded, quite ironically, that the most important Animal remains are now frequently used in groups in terms of taxonomic diversity and numerical archaeological ecological reconstructions and abundance were least studied within archaeology. Of interpretations. As indirect indicators of former course, archaeology puts the human species in the centre vegetations, and biotopes in general, the species of attention and animals that play(e d) a role in the food composition represented by the finds assemblages can provisioning of people or in other human economical be interpreted alongside more direct approaches within activities will therefore always receive disproportionally environmental archaeology, such as pollen analysis, the more attention. At the same time, however, it is clear study of seeds and fruits, and charcoal identification. that the information potential of many archaeological Experience revealed that carabid beetles (Carabidae) animal finds remains underexploited. form one of the most powerful groups amongst the After the initial, exploratory, studies ofa rchaeological ecological indicators in archaeology. Over the last insect remains, performed by multiple colleagues at the decades, fieldwork in Belgium has illustrated the 'Laboratory of Ecology' at Ghent University (Konjev validity of this statement (ERVYNCK eta!., 1996). Desender, Luc Mercken, Johan Mertens, Marc Pollet and Mark Van Kerckvoorde), it was Konjev Desender who solely continued this line of research, also after Carabid beetles in Belgian archaeology moving from Ghent University to the Royal Belgian Institute of Natural Sciences. Over the years, a growing Within Belgian archaeology, insects were used for the corpus of information was assembled, including material first time as ecological indicators in 1986, when remains from prehistoric sites (Bronze Age Kontich: ANNAERT et from sieved samples collected at two archaeological a!., 2004, Iron Age Zele: DESENDER & ERVYNCK, 2004), sites were handed over to researchers at the Laboratory Roman settlements (Braives: LENTACKER et al., 1993, for Ecology of Ghent University (then directed by J. Bruyelle: PIGIERE et al., 2001, 2002), Burst, Erpe-Mere: Huble). Ironically, the results of one of these earliest ERVYNCK et a!,. 1987, 1991, Elewijt: VAN IMPE et al., studies, dealing with material from an Iron Age site at 2005, Erps-Kwerps: LENT ACKER eta!., 1992, Liberchies: the 'Hogeweg' site, near Ghent (DESENDER, 1986e), are ScHELVIS et al., 2001, Merelbeke: DE CLERCQ et al., '' Carabid beetles and archaeology 37 THE NETHERLANDS e IS .16 • 19 FRANCE 0 40km >====== Fig. 1. - Location of the archaeological sites mentioned in the text (1: Antwerpen, 2: Braives, 3: Bruyelle, 4: Burst, 5: Elewijt, 6: Erps-Kwerps, 7: Hogeweg-Gent , 8: Kasterlee, 9: Kontich, 10: Liberchies, 11: Maldegem, 12: Mechelen, 13: Merelbeke, 14: Ravels, 15: Raversijde, 16: Sint-Andries, 17: Velzeke, 18: Zele, 19: Zerkegem). 2004, Ravels: VERHAERT et al., 2004, Sint-Andries, carabids against those of other insect groups (ERVYNCK Brugge: CoOREMANS et al., 2002, Velzeke: DESENDER & et al., 1994, 1996). In order to be useful as ecological ERVYNCK, unpublished data) and medieval occupations indicators within the archaeology of a certain region, an (Antwerpen: BUNGENEERS et al., 1989, Kasterlee: animal group should meet all of the following criteria: WOUTERS eta!., 1999, Raversijde: PIETERS eta/., 1999, (1) its remains must have a good chance to become Zerkegem: HoLLEVOET et al., 1994). The last study incorporated and survive in archaeological contexts, undertaken by Konjev Desender dealt with a sample (2) the remains should not be extremely rare, (3) they from the cesspit of the medieval prison of Malines must be identifiable to species level, ( 4) within the (Mechelen: LENT ACKER eta!., 2007). group, species variety must be high, (5) the ecology of The archaeo-entomological studies mentioned are the species involved must be sufficiently studied and characterised by a trend through time. Where the first tmderstood, (6) at least some of the species involved studies attempted to incorporate a wide variety of must have a limited ecological tolerance, and (7) the taxonomic groups, soon the focus became concenh·ated taphonomic history of the finds must be tmderstood upon carabid beetles (Carabidae ). This choice was and reliable. An evaluation of these criteria explains justified by comparing the characteristics of the why, for example, butterflies (Lepidoptera) are not very I I 38 A. ERVYNCK Main phyla Archaeozoologists rest rest Chordata Mollusca 1% 10% 3% Mollusca Arthropoda 2% Arthropoda 78% 85% Fig. 2. - Discrepancy between the number of species within different animal groups and the number of archaeozoologists studying their remains (after ScHELVIS, 1993). useful in archaeology (their remains typically do not present in a population, do not travel large distances, preserve in the soil), why dung beetles (Scarabaeoidea) implies that there is a causal relationship between are not very reliable (these flying insects are attracted their presence in an archaeological context and the over wide distances to archaeological structures, such enviromnent around it. as wells, and thus often bear no relationship with the Honesty necessitates underlining that, next to the local biotope(s) one wants to reconstruct), or why ecological considerations mentioned, focusing on flies (Diptera) are hardly ever used in archaeological carabids amongst the insect remains within Belgian ecological reconstructions (their archaeological enviromnental archaeology was also a pragmatic remains, often wing fragments, offer very limited choice. For some insect groups present in the sieved possibilities for identification). samples, specialists could (and can) simply not be In contrast, within the context of northwest found, or they show no interest in archaeological European archaeology, carabid beetles meet all of the material. Secondly, concentrating on a single, well criteria mentioned. The elements of their exoskeleton studied group, proved to be most effective, considering do survive in archaeological contexts and, when the balance between time spent and results obtained. present, are often found in considerable numbers. This approach, however, shows a stark contrast with Apparently, due to their walking instead of flying the development of the field in other countries. In the locomotion, they have always had a tendency to end Netherlands, for example, HAKBIJL (1989) explored up in an archaeological pitfall. For the specialist, with the full taxonomic potential of no iess than 27 families access to a good reference collection, identification of within the Coleoptera, in samples from an early Iron the head, thorax and elytra is often possible (Fig. 3). Age site in the Assendelver Polders. This tremendous Moreover, species variety is rather high in the region effort, however, has never been repeated. Within the considered and their ecology and distribution is well UK, again a wide variety of insect remains was studied, studied (DESENDER, 1986a-d, DESENDER et a/., 2008). both within a paleontological approach (as part of the Of course, the interpretation potential of a number of wider field of Quatemary palaeo-ecology, see ELIAS, eurytopic species, such as Abax ater, will always be 1994), or as a means to better understand the shucture limited compared to that of less tolerant species such and functioning of archaeological sites. Within the latter as, for example, Bembidion normannum, which only approach, defining indicator groups for anthropogenic occurs in salt marshes. In general, however, a sufficient deposits or human activities is vital, regardless of the nwnber of carabid species in the nm1hwest-Emopean insect or arthropod group to which they taxonomically fatma shows clear ecological characteristics, making belong (see e.g. CARROTT & KENWARD, 2001, and the them good ecological indicators. Finally, the fact that literatme there). The latter approach, ofhigh importance most carabid species, even when winged fonns are in archaeology, has never been followed in Belgium, Carabid beetles and archaeology 39 and the scarcity of material from the Metal (Bronze and Iron) Ages. However, this chronological loss of material and information does not help to understand the underrepresentation of material from the medieval and younger periods, versus that from Roman times. Possibly, the location of the sites investigated is partly responsible for this discrepancy. The Roman sites yielding carabid remains tend to be rural while the archaeology of the medieval and younger periods often concentrates on urban sites or large building complexes such as castles or abbeys. As carabids are less likely to occur in large numbers in these densely built-up areas, this could explain their relative scarcity in such sites. Indeed, the only medieval examples Fig. 3. - Remains of Carabus granulatus from a Roman well, excavated at Burst (see Fig. 1) (right), available derive from rural sites (Kasterlee: WouTERS compared to a recent reference specimen (left) eta!., 1999, Raversijde: PIETERS et a/.,1999, Zerkegem: (photo by the author). HoLLEVOET et a!., 1994) or from an urban site that was an open terrain at the moment of the deposition of the carabid remains (Antwerpen: BUNGENEERS eta!., 1989). Within the sample from the cesspit of the medieval mostly because of the lack of suitable, well-preserved prison ofMalines, only one carabid species was present archaeological contexts, and samples from them. (LENT ACKER eta/., 2007). On a smaller scale, preservation problems must be taken into account when evaluating different contexts Chronology and geographical distribution within an archaeological site. While the skeletal elements of the carabids are mechanically strong, Before attempting to evaluate the results of the and chitin, the most important building material, is archaeological analysis of carabid remains in Belgian chemically resistant against varying pH levels in the archaeology, some comments must be made explaining soil, the water regime of an archaeological context is the nature of the dataset available after more than of more importance. An alternation of dry and humid 20 years of (occasional) research. Firstly, it is clear conditions is detrimental while constantly moist or, that the data are unevenly spread through time, with even better, anoxic waterlogged conditions are ideal for an apparent bias towards Roman sites, and that the their survival. Carabid beetle remains can also survive northern part of the country is better represented than in constantly dry contexts (as in desert environments) other regions. Without doubt, these patterns are the but these tend not to occur in northwestern Europe. In result of varying excavation intensity, in itself the result general, the water regime explains why carabid remains of the different human impact (building activity) upon are mostly found in waterlogged archaeological the archaeological soil archives in different regions and contexts (wells, ditches, etc.) in Belgium. That carabid upon different types of sites. Moreover, the time and remains are most often preserved in wells can also effort spent on sieving sediment samples, determining help to explain why Roman material is more abundant the possibilities for the analysis of small animal than medieval finds. Wells are indeed more c01mnonly remains, varies widely between archaeological working excavated at Roman rural sites than in younger, urban groups. These characteristics can explain, for example, sites. Moreover, it could well be that Roman well why less carabid studies are available from Wallonia construction was better suited to serve as h·aps for compared to Flanders. Carabidae. There are indications that Roman wells, Additionally, the nature of the archaeological record in contrast to medieval examples, had no part above itself must also be considered. A recent survey of ground, making them more 'effective' as traps for archaeology in Flanders (see www.onderzoeksbalans.be) passing animals. shows that prehistoric sites, and certainly those with Recently, archaeological fieldwork in a prehistoric preserved organic remains, are relatively rare in the occupation zone at Lommel-Maatheide (see GEERTS northern half of Belgium. The erosion of time certainly et a!., 2006) revealed that beetle remains, including explains the absence of Stone Age carabid assemblages carabids, were present in meaningful numbers in peat '' 40 A. ERVYNCK deposits located off site. However, due to the very 4). This sedimentation process was interupted by three compacted and sticky nature of the peat, the carabid periods during which organic material could accumulate remains could not be extracted. Different methods to in the well. The carabid remains within these organic do so have been attempted but manual recovery seems layers show the succession of a beetle fauna from the only solution, a time consuming approach that has grasslands, over a species composition typical for not yet been followed. Still, this observation highlights barren grounds (fields), to an assemblage reflecting a the potential of such non-anthropogenic deposits for ruderal vegetation. The sequence again illustrates the the recovery of the Late Glacial and early Holocene phenomenon described earlier: Roman agricultural material that is now missing in the dataset. Possibly, overexploitation, i.e. ploughing of grasslands to turn where such peat layers have survived into the historical them into fields, causing soil destruction and rapid period, even much younger material could be recovered, abandonment of the site (ERVYNCK eta!., 1987). representing a natural, non-disturbed fauna that can be In general, the archaeological case studies compared against the archaeological assemblages. In show that the impact of people was already severe, this way, archaeological carabid research in Belgium centmies before the Industrial Era, quite certainly as could finally make a link with the paleo-ecological early as Roman times. This was clearly the case for work on Quaternary insects, which has received much (local) deforestation, reflected by a general scarcity attention in neighbouring countries (see ELIAS, 1994 for of stenotopic forest species amongst the carabids a review). excavated. At the same time, however, it should not be concluded that untouched environments no Archaeology and presentday ecology When the results of the carabid samples from Belgian archaeological contexts (see the publications cited above) are put together, the ecological information gained, although of vital importance for the understanding of the sites, could be evaluated as being mostly of an anecdotal nature, and of little direct use for studies of present-day ecology. However, while it is true that each site rep01i has to be seen as a single sample, and that much more data are needed, it has been possible to arrive at conclusions that have a more general application. This was possible by the incorporation of the ecological reconstructions within much broader frameworks of former human behaviour. The data for a Roman site at Merelbeke, for example, indicating the presence of fields in a landscape where on the basis of the soil characteristics only grasslands would be expected (DE CLERCQ et a!., 2004), fit into the general idea of a large-scale Roman agriculture characterised by the overexploitation of poor soils, exemplified by turning them from pastures into fields. All over the Low Countries, subsequent exhaustion of the soil, and erosion, soon ended these unfortunate agricultmal projects (GROENMAN -VAN WAATERINGE,l983). Interpretations become much more solid, of comse, when a succession of faunas can be studied from a single structure. A rare opportunity for doing so was provided by the excavation of a Roman well at Burst, near Aalst. The fill of this structure showed that, at Fig. 4. - Stratigraphy of the fill of a Roman well, excavated multiple occasions, sediment was deposited into the at Burst (see Fig. 1) (photo M. Pieters, Flemish shaft and sorted into layers of sand, silt and clay (Fig. Heritage Institute). Carabid beetles and archaeology 41 0 woodland 0 eurytopic 0 shrubs • open land II 10 9 8 7 ~ >-. 6 ..:s 5 4 3 2 0 20 40 60 80 100 %species Fig. 5. - Sequence of the most important ecological groups of Carabidae in the fill of a Roman well at Velzeke (see Fig. 1) (Desender & Ervynck unpublished data). longer occured in Roman times. This is, for example, Velzeke has put an extra emphasis upon the fact that illustrated by an unpublished sequence of material from not only the cultural or historical interpretations are a Roman well excavated near a temple site at Velzeke important but that the data are also relevant for the (near Zottegem) (RoGGE, 1988), deposited from the late study of the present ecology of carabid faunas and the Roman to the early medieval period (dates on the basis environments in which they occur. Indeed, the processes of radiocarbon measurements) (Fig. 5). In the lowest documented by environmental archaeology have part of the fill, a carabid fauna was found dominated by defined the composition of the floras and faunas we species of forest and shrubbery, with an admixture of find now, and this is particularly true for Carabidae. The species of open vegetation (cultural land). This suggests poor mobility and colonisation capacity of most species that the well was located in a forested environment, not within this group implies that (even sliort) changes in an unusual situation for a temple site. Higher up in the an environment provoke long-lasting alterations in the sequence, the species of open vegetation disappear, carabid species composition (see DESENDER & VANDEN without doubt reflecting the abandonment of the Roman BussCHE, 1998; DESENDER eta!., 1999). Consequently, site, after which forest regained the terrain. In the early from a time perspective, the actual carabid faunas medieval deposits, species indicating open vegetation provide an archive from which the impact of humans in reappear, the result of the wide-scale recolonisation of an often-distant past can be infened. lands in that period (DESENDER & ERVYNCK, unpublished In general, the present state of the research into data). The changing frequency of forest species in the archaeological and actual carabid faunas seems to carabid sequence thus documents a broad demographic allow an integration of the datasets, stimulating a trend in the human population of Flanders, but proves, at broader time perspective for ecological studies. The the same time, that forest never disappeared completely archaeological data make it clear that 19th century during Roman times. scientific infonnation or old museum collections are Within the development of the study of not sufficient as references when reconstructing so archaeological carabid remains, the case study from called 'original' faunas (not significantly influenced by II 42 A. ERVYNCK human impact). The already mentioned Roman carabid vroege IJzertijd op de site Kapelleveld in Kontich (provincie collection from Velzeke contained a richer species Antwerpen). Archeologie in Vlaanderen, VIII: 79-103. spectrum of stenotopic forest dwellers than has ever BucKLAND, P. C., 1976. The use of insect remains in the been found in an actual forest environment in Belgium. interpretation of archaeological environments. In: DAVIDSON, This means that even the most 'natural looking' present D. A. & SHACKLEY, M. L. (Editors), Geoarchaeology: Earth day forests have been heavily influenced by human sciences and the past. Duckworth, London, pp. 360-396. activities and, in terms of carabid fauna, no longer BUNGENEERS, J., DESENDER, K. & ERVYNCK, A., 1989. represent an original situation (DESENDER eta!., 1999). Keverresten uit de waterput. Scharnier (Antwe1pen), 6: 5-7. Inevitably, the integration of archaeological and CARRoTT, J. & KENWARD, H., 2001. Species associations among actual data promoted a new methodological approach. insect remains from urban archaeological deposits and their Instead of comparing actual biotopes with the significance in reconstructing the past human environment. archaeological reconstructions of former environments Journal ofA rchaeological Science, 28: 887-905. (largely based upon a subjective evaluation of the CoOPE, G. R., 1978. Constancy of insect species versus presence, and sometimes abundance of species), a more inconstancy of Quaternary environments. In: MoUND, L. A. integrated approach necessitated the incorporation & WALOFF, N. (Editors), Diversity of insect faunas. Royal of the basic archaeological identifications into Entomological Society of London Symposium 9. Royal databases of recent biodiversity surveys. Doing this, Entomological Society, London, pp. 176-187. the archaeological assemblage is treated as any other COOREMANS, B., DESENDER, K., ERVYNCK, A. & SCHELVIS, J., sample and comparisons are made by statistical analyses 2002. Onderzoek van plantaardige en dierlijke resten uit (see DESENDER eta!., 1999 for forest species). een Romeinse waterput van de vindplaats 'Refuge' te Sint Andries, Brugge (provincie West-Vlaanderen): economie en ecologie. Archeologie in Vlaanderen, VI: 209-229. Future prospects DE CLERCQ, W., BASTIAENS, J., DEFORCE, K., DESENDER, K., ERVYNCK, A., GELORJNI, V., HANECA, K., LANGOHR, R. & VAN At the moment of the untimely death of Konjev PETEGHEM, A., 2004. Waarderend en preventief archeologisch Desender, archaeological carabid research in Belgium onderzoek op de Axxes-locatie in Merelbeke (provincie had a bright future, certainly also through the Oost-Vlaanderen): een grafueuvel uit de bronstijd en een integration into actual ecological research frameworks. nederzetting uit de Romeinse periode. Archeologie in Moreover, the integrated approach made it easier to Vlaanderen, VIII: 123-164. present the research results to a wider public (see e.g. DESENDER, K., 1986a-d. Distribution and ecology of Carabid DESENDER 2009, published posthumously). As the years beetles in Belgium (Coleoptera, Carabidae). Part 1-4. of data collecting are now beginning to produce higher Koninklijk Belgisch Instituut voor Nah1urwetenschappen, interpretation levels, Konjev's research must certainly Brussel, 30, 24, 23 & 48 pp. be continued. New specialists will have to be convinced DESENDER, K.,1986e. Hogeweg. Keverfragmenten to work with archaeological material and, in a changing in sedimenten uit de Vroege IJzertijd (unpublished world in which archaeology becomes more and more manuscript). commercialised, means will have to be sought to allow this to happen. Otherwise, archaeological carabid DESENDER, K., 2009. Miskende getuigen: loopkevers in het Meerdaalwoud. In: BAETE, H., DE BrE, M., HERMY, M. & VAN research in Belgium will only have known a promising DEN BREMT, P. (Editors), Mirada!. Erfgoed in Heverleebos en start. .. Meerdaalwoud. Davidsfonds, Leuven, pp. 217-233. DESENDER, K., DEKON!NCK, W. & MAES, D., 2008. Een nieuwe Acknowledgements verspreidingsatlas van de loopkevers en zandloopkevers (Carabidae) in Belgie. lnstituut voor Natuur- en Many thanks go to Sheila Hamilton-Dyer for the correction of the Bosonderzoek, Brussel, 184 pp. English, and to Alexis Wielemans (Flemish Heritage Institute) for DESENDER, K. & ERVYNCK, A., 2004. 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