Abstract Abstract
‘Carabidologists do it all’ (Niemelä 1996a) is a phrase with which most European carabidologists are familiar. Indeed, during the last half a century, professional and amateur entomologists have contributed enormously to our understanding of the basic biology of carabid beetles. The success of the field is in no small part due to regular European Carabidologists’ Meetings, which started in 1969 in Wijster, the Netherlands, with the 14th meeting again held in the Netherlands in 2009, celebrating the 40th anniversary of the first meeting and 50 years of long-term research in the Dwingelderveld. This paper offers a subjective summary of some of the major developments in carabidology since the 1960s. Taxonomy of the family Carabidae is now reasonably established, and the application of modern taxonomic tools has brought up several surprises like elsewhere in the animal kingdom. Progress has been made on the ultimate and proximate factors of seasonality and timing of reproduction, which only exceptionally show non-seasonality. Triggers can be linked to evolutionary events and plausibly explained by the “taxon cycle” theory. Fairly little is still known about certain feeding preferences, including granivory and ants, as well as unique life history strategies, such as ectoparasitism and predation on higher taxa. The study of carabids has been instrumental in developing metapopulation theory (even if it was termed differently). Dispersal is one of the areas intensively studied, and results show an intricate interaction between walking and flying as the major mechanisms. The ecological study of carabids is still hampered by some unresolved questions about sampling and data evaluation. It is recognised that knowledge is uneven, especially concerning larvae and species in tropical areas. By their abundance and wide distribution, carabid beetles can be useful in population studies, bioindication, conservation biology and landscape ecology. Indeed, 40 years of carabidological research have provided so much data and insights, that among insects - and arguably most other terrestrial organisms - carabid beetles are one of the most worthwhile model groups for biological studies.
Keywords: Carabidae, ground beetle, systematics, biology, life history, rhythms, seed feeding, ant feeding, ectoparasitism, predation on amphibians, dispersal, pitfall trapping, statistics, population dynamics, long-term research, bioindicators, conservation, habitat management, landscape ecology
1 Introduction
1.1. General
Carabid beetles are one of the best-known taxa in entomology. These beetles have been studied intensively by generations of coleopterists, who have clarified the taxonomy and phylogeny, geographic distribution, habitat associations and ecological requirements, life history strategies and adaptations, especially in Europe (e.g. Holdhaus and Lindroth 1939; Palmén 1944; Lindroth 1945a, b, 1949; Thiele 1977; Ball 1979; Desender 1986, Desender et al. 1994a; Turin 2000; Luff 2007).
This wealth of basic information has fostered a plethora of quantitative ecological studies. Indeed, the first European Carabidologists’ Meeting in Wijster, the Netherlands in 1969, touched upon one of the fascinating characteristics of carabid beetles – dispersal and dispersal power (Den Boer 1971). As a life history trait, dispersal has profound consequences for the dynamics and persistence of populations, the distribution and abundance of species and for community structure (Dieckmann et al. 1999). Not surprisingly, a summary based on the 3rd International Carabidologists’ Meeting emphasised the role of dispersal in increasingly fragmented landscapes, and argued that much more knowledge on the effects of habitat loss and fragmentation on carabid beetle population dynamics is needed if sensible decisions are to be made regarding conservation and land-use (Thacker 1996).
But why study carabid beetles? The reasons are diverse: relatively stable taxonomy, high species richness, occurrence in most terrestrial environments and geographical areas, the availability of easy collection methods, known sensitivity to environmental changes, and perceived role as beneficial in agriculture (see Darlington 1943; Lövei and Sunderland 1996; Rainio and Niemelä 2003). Armed with such a diverse wealth of knowledge, many ecologists and taxonomists have turned to carabid beetles to test ecological research questions. In this paper we emphasise progress in some of the major fields in carabidology since the first European Carabidologists’ Meeting, 40 years ago.
1.2. Basic knowledge
Modern disciplines in carabid beetle ecology, such as bioindication, conservation and habitat management, landscape ecology and urban ecology rely heavily on the work done by professional and amateur carabidologists from the more traditional fields of natural history, systematics and taxonomy. This species-rich family occurs in most terrestrial habitats and is found in the vegetation as well as high up in the trees and the canopy, not only in the tropics (Arndt 2005). This is probably the main reason why carabids are relatively well represented in collections around the world. In many regions, information on labels from these collections has been gathered in large databases. Combined with data from systematic sampling, such datasets enable profound faunistic work. These databases are increasingly elaborated and published as annotated checklists, red lists, catalogues and/or atlases. In combination with a clear taxonomy, mainly identification literature, these provide a sound basis for biogeographical, biological, ecological and experimental studies. Table 1 shows an overview of the major publications for the European continent, which is covered well, although there is clearly need for updating in a few regions, mainly in the east (Romania, Hungary, Russia, Caucasus). In some cases, older works are mentioned in Table 1, which belong to antiquity and do not adequately cover the fauna of that region anymore (e.g. Ganglbauer 1892; Apfelbeck 1904; Porta 1923–1959). These older works are hardly in use for identification anymore. However, they still provide historical bases for modern identification works, which often have to be elaborated from numerous smaller keys or large revisions (e.g. Jeannel 1926–28; Breuning 1932–37), such as the keys to the Carabinae (Casale et al. 1982) and to the supra-specific taxa of Italy (Casale 2005).
Table 1.
Overview of publications concerning the faunistics of ground beetles in Europe.
A sound basic list of the Carabidae of the world is the recent checklist published by Lorenz (2005) and a catalogue with distributional data is available for the Palaearctic region as a whole (Löbl and Smetana 2003). Furthermore, many recent checklists and catalogues are available (concerning Europe, see some examples in Table 1). In particular, Kryzhanovskij et al. (1995) provided detailed information on the carabid fauna of Russia and adjacent countries (including central-Asiatic). In the Western Hemisphere (the Americas), detailed information is available, especially for the regions north of Mexico (Lindroth 1961–1969; Ball and Bousquet 2001; Larochelle and Larivière 2003; Erwin 2007; Erwin and Pearson 2008), or will soon be (Erwin in preparation), but in many tropical areas of Central and South America, many genera and species remain undescribed. Other geographical areas are less well known. Asia, as a huge continent is relatively well-known in some parts, such as Siberia, Near and Middle East and especially Japan (e.g. Habu 1967, 1973, 1978), whereas immense areas are a “work in progress” (China, The Himalayas and South-East Asia). Africa is well-known in some northern countries, in particular Morocco, Algeria and Tunisia, thanks to the contributions of specialists like Antoine (1955–1962), Bedel (1899–1900) and Kocher (1963). Nevertheless, in spite of the numerous papers published by Alluaud, Basilewsky, Jeannel and others, the sub-Saharan (tropical) part of the continent needs more investigation. Australia, thanks to the C.S.I.R.O. has one of the best-organised services of insect collections, and is covered by catalogues and revisions, of which we highlight the catalogue by Lawrence et al. (1987). But also, recent investigations allowed the discovery of many new genera and species, including impressive, large sized Pamborus species.
Finally, remote islands and archipelagos such as like Madagascar, Papua-New Guinea and Galápagos, for instance, have been carefully investigated by specialists like Jeannel, Darlington and Desender, respectively, but produce many new discoveries every year.
In the world catalogues of (Lorenz (1998, 2005) more than 35 000 ground beetle species have been listed. An estimated number of 40 000 species, which is more than 10 times the number of described mammals, has often been mentioned (Thiele 1977; Noonan 1985). Currently, approximately 38 600 valid names occur worldwide (based on Lorenz 2005 and an estimate of approximately 100 additional new species every year). For the Western Hemisphere only, the species count currently stands at 9 374 (Terry Erwin in litt.).
More in line with the meetings are a number of thematic treatments, but again the listed works are only examples. For a more complete and thematically arranged overview of significant work in carabidology, we refer to the excellent introduction to the proceedings of the Symposium on Phylogeny and Classification of Caraboidea by Ball et al. (1998). Worth mentioning for European carabidology are the publications of the German “Gesellschaft für Angewandte Carabidologie” (GAC) with special reference to habitat studies, such as carabid beetles in river meadow habitats (GAC 1999), in forests (GAC 2001) and in xerothermic habitats (GAC 2004). The GAC provides many carabidological papers in open access (see http://www.laufkaefer.de/gac). Other published thematic studies, often including compilations of numerous papers from various authors, concern, amongst others: biotopes (Heydemann 1962; Schjøtz-Christensen 1965), larvae (Brandmayr and Zetto Brandmayr 1982; Arndt 1991; Luff 1993), biology and periodicity (Larsson 1939), agroecology (Holland 2002), biogeography (Ball 1985; Noonan et al. 1992), dispersal ecology (Palmén 1944; Den Boer 1977; Baars 1982; Desender 1989b; Aukema 1995), morphology (Sharova 1981; Deuve 1993) and phylogeny (Ball et al. 1998). This listing is not exhaustive, especially in the fields of genetics and molecular biology, which are growing rapidly. We conclude with the classical works Die Fennoskandischen Carabidae (Lindroth 1945a, b, 1949, re-published in English as Lindroth 1988, 1992a, b) and Carabid beetles in their environments (Thiele 1977). These inspired many carabidologists and have been, for many students, the starting point of their enthusiasm.
1.3. European Carabidologists’ Meetings (ECMs)
In 1959, Piet den Boer, a zoologist at the Biological Station in Wijster, started pitfall trapping at several locations in the Dwingelderveld, a large area of heathland. His purpose was to test the model proposed by Andrewartha and Birch (1954), in which animal populations could be thought of as sets of smaller local populations which periodically become extinct, their sites being subsequently reoccupied. This became known (and fashionable) under the term “metapopulation” (Levins 1970). By using carabid beetles as test organisms, Den Boer was able to show that in a large area many local populations or interacting groups fluctuate in numbers of individuals in space and time, developing his theory of ‘spreading of risk’ (Den Boer 1968). According to this theory, species occupying large areas survive more easily because the reproductive success of each separate (but interacting) group differs at different places. Dispersal between these interacting groups stabilises the number of individuals in the whole population through time. Local extinctions may occur but the chances of extinction of the entire population are minimised (Den Boer 1970). Den Boer eagerly wanted to discuss this topic with other carabid beetle specialists, in particular with Carl Lindroth from Sweden, who studied the significance of dispersal and Hans-Ulrich Thiele from Germany, who studied the reproduction of these animals. Consequently in 1969, a number of eminent European carabidologists were invited to Wijster. This select group of researchers focused on the topic of dispersal and the dispersal power of carabid beetles (Fig. 1a). In 1973, Thiele invited a number of carabidologists to Rees-Grietherbush, a field station of the University of Cologne. This second ECM appeared to be an informal one and no proceedings volume was published. However, it resulted in the organisation of a now official third ECM, also at Rees-Grietherbush, by Thiele and his colleague Friedrich Weber in 1978. Most participants were German or Dutch, though Pietro Brandmayr from Italy was also present. The proceedings entitled ‘On the evolution and behaviour of carabid beetles’ was dedicated to Lindroth, who passed away in early 1979. In 1981, Weber took the initiative and organised the fourth ECM at Haus Rothenberge (Münster), on the theme ‘The synthesis of field study and laboratory experiments’. Thiele presented a lecture but his contribution for the proceedings was never received. The proceedings, dedicated to Thiele, was published after his death in 1983.
Figure 1a.
Participants of the first European Carabidologist Meeting in Wijster, 1969. From left to right: Vlijm, Van der Aart, Lindroth, Stein, Wijmans, Hengeveld, Palmén, Van Dijk, Richter, Venema, Mook, Thiele, Tjallingii, Den Boer, Haeck, Neumann, Meijer.
The first four meetings were followed by meetings organised across Europe (Table 2). As a result of political changes in Eastern Europe since the 1990s, the ECMs attained a more ‘complete’ European character. Not only did it become easier for scientists from Eastern Europe to attend these meetings, they also started to organise them. Even more noticeably during recent decades, carabidologists from beyond Europe regularly started to participate in the ECMs. Besides the official ECMs, there have been a few separate carabid beetle meetings in Europe (Table 2). Two of these (Hamburg in 1984 and Kauniainen in 1995) were not official ECM meetings, though they were mainly attended by the same carabidologists who regularly attend ECMs. The fourteen proceedings from the major ground beetle meetings that have been published before the present volume (see Fig. 1b-c, Table 2), comprise together more than 400 articles covering a wide range of topics. A rough classification of the articles leads to the following summary: Habitat preference, community ecology was the topic of 84 papers, Biology (development, preferences, etc.) of 55, Population biology - 46, Nature conservation - 35, Agro-ecology - 34, Dispersal ecology - 33, Evolutionary biology, phylogeny - 22, Morphology - 15, Ecology, general - 13, Genetics - 13, Biogeography - 11, Taxonomy - 11, Method-development - 10, Rest – 10, Faunistics - 9, and Palaeontology - 2. A similar series of meetings and proceedings started in America with the publication of the First International Symposium of Carabidology (Erwin et al. 1979). In 1999, a volume consisting mainly of taxonomic papers was published, dedicated to the memory of Oleg L. Kryzhanovskij (Zamotailov and Sciaky 1999).
Table 2.
The year, location, title and editors of all the European Carabidologists’ Meetings.
| Year | Location | Proceedings |
|---|---|---|
| 1969 | Wijster, The Netherlands (ECM 1) | 1971. Dispersal and dispersal power of carabid beetles (Den Boer) |
| 1973 | Rees-Grietherbush, Germany (ECM 2) | None |
| 1978 | Rees-Grietherbush, Germany (ECM 3) | 1979. On the evolution of behaviour in carabid beetles (Den Boer et al.) |
| 1981 | Münster, Germany (ECM 4) | 1983. The synthesis of field study and laboratory experiments (Brandmayr et al.) |
| 1982 | Stara Brda Pilska, Poland(ECM 5) | 1986a. Feeding behaviour and accessibility of food for carabid beetles (Den Boer et al.) |
| 1984 | Hamburg, Germany (17th International Entomological Congress) | 1986b. Carabid beetles, their adaptations and dynamics (Den Boer et al.) |
| 1986 | Balatonalmadi, Hungary (ECM 6) | 1987. Proceedings of the 6th ECM (Den Boer et al.) |
| 1989 | London, United Kingdom (ECM 7) | 1990. The role of ground beetles in ecological and environmental studies (Stork) |
| 1992 | Louvain la Neuve, Belgium (ECM 8) | 1994a. Carabid beetles, ecology and evolution (Desender et al.) |
| 1995 | Kauniainen, Finland (3rd International Carabidology Congress) | 1996b. Population biology and conservation of carabid beetles (Niemelä) |
| 1998 | Camigliatello, Italy (ECM 9) | 2000. Natural history and applied ecology of carabid beetles (Brandmayr et al.) |
| 2001 | Tuczno, Poland (ECM 10) | 2002. How to protect or what we know about carabid beetles (Szyszko et al.) |
| 2003 | Århus, Denmark (ECM 11) | 2005. European Carabidology 2003 (Lövei and Toft) |
| 2005 | Murcia, Spain (ECM 12) | 2006. Proceedings of the XII ECM; ground beetles as a key group for biodiversity conservation studies in Europe (Serrano et al.) |
| 2007 | Blagoevgrad, Bulgaria (ECM 13) | 2008. Back to the roots and back to the future. Towards a new synthesis between taxonomic, ecological and biogeographical approaches in carabidology (Penev et al.) |
| 2009 | Westerbork, Netherlands (ECM 14) | 2011. Present volume (Kotze et al.) |
Figure 1b.
Front covers of the first European meetings, ECM 1–8 and that of Hamburg 1984 (centre cover) (see also Table 2).
Figure 1c.
Front covers of the last five ECMs and of a few major carabidology publications (Thiele 1977; Ball et al. 1998; Erwin et al. 1979; Noonan et al. 1992) (see also Table 2).
In 2009, the 14th ECM returned to the starting grounds in the Netherlands and was attended by five participants of the first ECM: Piet den Boer, Jaap Haeck, Rob Hengeveld, Jan Meijer and Theo van Dijk. The participants visited the permanent sampling plots in the Dwingelderveld and Mantingerveld, started 50 years earlier.
2 Systematics, phylogeny and evolution
2.1. Overview
Regular carabidologists’ meetings have contributed significantly to our understanding of carabid phylogeny, evolution and systematics, as evidenced by the presentation of more than 60 papers on these topics. Progress has been made at different taxonomic ranks and in different fields of carabid systematics. At present, the integrative approach of combining morphology, molecular systematics, ethology, ecology, geographic distribution, etc., as well as the use of bioinformatics, is recognised as the best framework for solving the challenges still faced by carabidologists (Assmann et al. 2008), and by animal taxonomists in general.
What follows is a short overview of recent advances in carabid beetle systematics, concentrating on literature presented at ECMs and the international congresses mentioned above. As the main aim of this section is to present a general overview, only some of the main papers with a wide scope are cited.
2.2. General outline on systematics and phylogeny of the Carabidae
Ball (1979) showed that the classification of Carabidae is mostly based on morphological characters and that it includes both clade-based and grade-based criteria; classifications differ depending on the importance given to one or the other criterion. After this seminal revision, few advances have been made to unify the criteria to elect Caraboidea (splitters) or Carabidae (lumpers), and the same holds true for other high-ranked taxa. A practical synthesis of these ideas was presented by Nagel (1979a), while Ball et al. (1998) and Assmann et al. (2008) revised the issue in depth. These two last-mentioned papers highlighted the need for an integrative approach to morphology, morphometrics and molecular systematics as the appropriate way of finding rapid solutions for challenging problems.
2.3. Within-species diversity
An electrophoretic study on 14 Pyrenean populations of Carabus punctatoauratus (Assmann 1990) revealed that the Pyrenees probably hosts an isolated relict population for this species, and that bottlenecks have affected western, central and eastern populations differentially. Subtle differences at a micro-geographic scale have also been shaped by small bottleneck phenomena in this species with low dispersal power.
Range expansion of Carabus auronitens during the 19th century has allowed gene flow between populations in the surroundings of Münster, Germany, as evidenced by an electrophoretic study of 19 populations that showed a steep gradient of slow and fast alleles (Terlutter 1990). The high dispersal power of this species accounts for the observed allelic gradient (esterase-encoding gene) from source areas to recently colonised areas (Niehues et al. 1996). Assmann et al. (1994) showed that present-day populations of this species originated from three major refuges in southern France and that these putative core populations have contributed differentially to postglacial range expansion of the species.
Ashworth (1996) showed that Quaternary climatic oscillations did not lead to enhanced rates of extinction and speciation in carabids, as inferred from 14C-dated fossil assemblages. The future responses of Carabidae to climate change will probably be similar to that of the past, with the exception that extinction rates are expected to be higher because of human-caused habitat fragmentation.
Rasplus et al. (2000) found that populations of the threatened species Carabus solieri consist of two distinct clusters corresponding to subspecies bonnetianus and solieri. These populations were probably isolated during the last glaciation and are worthy of protection as gene flow is restricted between these two groups. Moreover, molecular markers suggest that the subspecies curtii is a hybrid between bonnetianus and solieri.
Desender et al. (2000) investigated the genetic diversity and wing polymorphism of the salt-marsh beetle Pogonus chalceus in 30 populations from the Atlantic coast and nine populations from the Mediterranean Basin. These Mediterranean populations showed little differentiation associated with high dispersal power, a finding possibly related to habitat instability. A higher structuring was found in Atlantic populations, which showed varying degrees of wing polymorphism and dispersal power, possibly related to adaptation to particular conditions.
Kamer et al. (2008) investigated variation in the 12S RNA sequence in populations at different geographic scales, namely the Baltic coast, inland populations across Central Europe, and Central plus Western Europe. Population structure varied as a result of complex factors that include past history and present dispersal power, amongst others. Cryptic taxa or a lack of molecular differences among siblings were also found, showing the usefulness of landscape genetic analyses.
2.4. Species borders and hybridisation
Koch (1986) showed that Pterostichus nigrita and its sibling Pterostichus rhaeticus are distinct species according to habitat preferences, subtle details in male and female genitalia and karyotypic numbers. Both species are reproductively isolated, as shown by crossbreeding laboratory experiments. More recently, Angus et al. (2008) described a new cryptic species in the Iberian Peninsula, Pterostichus carri, and a new subspecies of Pterostichus nigrita from Anatolia. All taxa shared a basic 2n = 36 + X male karyotype, whereas marked variation in the number of accessory chromosomes was found within and between these taxa.
Vogler and DeSalle (1994) analysed the relationships of 17 populations of Cicindela dorsalis along a littoral transect from New England to Veracruz. These populations are currently ascribed to four subspecies which is difficult to ascertain. Mitochondrial DNA haplotypes showed that populations could readily be grouped into two major entities that represent well defined phylogenetic species without gene flow between them, one occupying the Atlantic coast, the other inhabiting the Gulf of Mexico. Within each of these entities, moderate diversification was found but without much geographic structure, probably because of moderate gene flow between populations.
Galián et al. (1996) studied the karyotypes and the RFLPs resulting from digestion of total DNA with endonuclease EcoRI in four populations ascribed to Ceroglossus chilensis. Differences between these populations in terms of chromosome number and molecular data led to the conclusion that there are three cryptic species living in sympatry.
A clear distinction between Abax parallelepipedus and Abax angustatus (reported as a subspecies of the former) resulted from a morphological analysis of sympatric populations of both species, and a molecular study based on allozymes and mitochondrial DNA (Düring 2002). No molecular evidence of hybridisation between these two species was found.
Mossakowski et al. (1986) carried out a field study on the frequency of hybrids between species of the subgenus Chrysocarabus, Chrysocarabus lineatus and Chrysocarabus splendens in the Pyrenees. Reliable morphological characters allowed for determining the occurrence of hybrids. Both species may hybridise (up to 40% of individuals) when particular ecological conditions are met, which indicates that complete reproductive isolation has not yet been attained. However, a number of characters are fixed in each species allowing their classification as valid species. Furthermore, Düring et al. (2000, 2006) studied the mitochondrial haplotype in many Chrysocarabus splendens populations and found convincing evidence of introgressive hybridisation in Chrysocarabus (incongruence between mitochondrial and nuclear gene trees). In contrast, nuclear ITS-2 sequences showed that populations of Chrysocarabus splendens made up a monophyletic clade, which is sister to that made up by Chrysocarabus lineatus and Chrysocarabus lateralis. Shared haplotypes between Chrysocarabus splendens and Chrysocarabus punctatoauratus are probably the result of introgression of the latter into the former species. On the other hand, mitochondrial DNA of Chrysocarabus rutilans was probably acquired from Chrysocarabus splendens through introgression.
2.5. Speciation, radiation and biogeography
Juberthie (1979) analysed the evolutionary pathways of the genus Aphaenops (Trechinae) from putative epigean ancestors to specialised troglobionts, and noted that food must have been a major factor in promoting their morpho-functional characters. He also concluded that Aphaenops and other hypogean Trechinae are not living fossils but show highly derived characters, either regressive (loss of eyes and pigmentation) or positive (slender appendages, new chemoreceptors) with regard to ancestral epigean forms, with which they still share particular plesiomorphies.
Mossakowski (1979) postulated that habitat preference is an evolutionary process that can be reconstructed when matching it against a phylogenetic tree of particular taxa. He tested this hypothesis by considering the subgenus Chrysocarabus and concluded that there was an adaptive shift from Mediterranean to deciduous forests and a recent colonisation of alpine environments.
Liebherr (1986) constructed a phylogeny of the Agonum extensicolle group based on morphological quantitative characters and the allelic frequencies derived from the electrophoresis of soluble enzymes. The resulting tree was used to test the hypothesis of the vicariance effects of the Cochise filter/barrier separating the Sonoran and Chihuahuan deserts in SW North America. He argued that the zone between the deserts has probably caused vicariant events between particular pairs of species and species groups, and also between subspecies of Agonum decorum 6.5 to 2.8 million years ago. This barrier has probably led to the same phenomena in other carabid taxa.
Desender et al. (1990) studied speciation of the genus Pterostichus in the Galápagos using multivariate morphometric analysis and ecological data. They concluded that a combination of allopatric (stepping stone model) and parapatric events (segregation in altitude of two species inhabiting the same island) may explain radiation of the genus from ancestors related to Pterostichus peruviana, a species presently found in South America.
Andersen and Skorping (1990) presented a conclusive model of sympatric speciation of the genus Bembidion (and in particular in the subgenus Chysobracteon), in which habitat selection and the effects of parasites may give rise to disruptive selection that promotes reproductive isolation and in turn speciation. Habitat shifts in riparian carabids may have evolved in sympatry, whereas allopatry would have produced new taxa showing mere variations of the same ecological theme.
Baehr (1994) constructed a cladistic analysis of the Pseudomorphinae based on morphological characters that solved relationships of the main lineages within the subfamily. He postulated that the subfamily has an Australian-South American origin, and that it has recently spread to North America and SE Asia.
Brandmayr and Zetto Brandmayr (1994) presented an elaborated hypothesis on the evolutionary history of the genus Abax, based on characters of male genitalia (inflated median lobe), larval morphology, type of parental care and larval behaviour, habitat preferences and geographic distribution. Ancestors of this genus possibly inhabited lowland forests during the late Miocene, whereas most recent taxa are found in alpine grasslands and mountain forests. This suggests that there has been a major colonisation trend towards mountains during the last geological periods. A predominantly allopatric pattern was inferred for the radiation of Abax.
The supertribe Carabitae poses major evolutionary problems because many character states are difficult to interpret due to homoplasy, and the biogeographic patterns of tribes are not congruent at first glance with relationships derived from molecular and morphological data. A synthesis of different studies (Prüser and Mossakowski 1998; Kamer et al. 2002; Mossakowski 2002) based on the analysis of morphological characters (adults and larvae) plus molecular data, indicates that Cychrini is sister to all other tribes, and that Carabini is sister to a clade made up of tribes Ceroglossini and Pamborini. This hypothesis also postulates a Laurasian origin of Carabitae and a single migration event across the tropics. A corollary of this hypothesis is that the Cychrus-like mandible of Pamborus is a homoplasy that would result from an adaptation to feed on snails (‘cychrisation’).
Of the four Calosoma species inhabiting the Galápagos, only Calosoma granatense is widespread among islands and altitudinal habitats. In spite of its high dispersal power and morphological stability, this species shows substantial genetic differentiation between populations on different islands and volcanoes (Desender and Verdyck 2000). There was probably a single colonisation event from the mainland and a stepping-stone model of island colonisation. However, gene flow must have been enough to prevent speciation events. The other three Calosoma species of the Galápagos are endemic to localities at high altitudes on a single island, which suggests that they have originated by convergent habitat shifts.
The phylogenetic relationship of three Carabus species inhabiting the Tenerife and Gran Canaria (subgenus Nesaeocarabus) was investigated by a phylogenetic analysis based of the mitochondrial nd5 gene (Prüser et al. 2000). The hypothesis of a close relationship between Nesaeocarabus and the subgenus Eucarabus was rejected. Instead, Canarian taxa were closely related to the subgenus Eurycarabus from northern Africa, southern Italy, Sardinia and Sicily. Diversification of Nesaeocarabus in the Canaries was congruent with the geological history of the archipelago, with a diversification of ancestors beginning 14–7 million years ago.
The subgenus Platycarabus includes five species living in the Alps and adjacent areas. Casale et al. (1998) tested the hypothesis of a close relationship of these species with the subgenus Hygrocarabus, both included in the genus Chaetocarabus sensu Ishikawa (1984). Separate and combined analyses of 26 adult and larval characters, and of sequences of the nd1 gene, rejected this hypothesis, as Platycarabus is a robust monophyletic lineage distantly related to Chaetocarabus, and is even farther from Hygrocarabus.
Mossakowski (2005) revised the proposal of Imura (2002) of grouping the genus Carabus s. l. into 29 sections and 137 genera, based on molecular data (see also Casale and Mossakowski 2003). Analysis of the inflated median lobe of the male endophallus and the reassessment of DNA sets with stringent criteria of bootstrap values showed that (i) relationships of the subgenera of Carabus were poorly solved, (ii) the results do not support the hypothesis of an explosive radiation of the ancestors of this genus, and (iii) these uncertainties do not favour the ranking of subgenera to genera proposed by Imura (2002).
2.6. Phylogeny based on different types of characters
Ethology
Brandmayr and Zetto Brandmayr (1979) found that the genus Abax shows different stages between a pure pre-social condition of merely laying eggs with a well-developed ovipositor, and the advanced construction of a chamber, laying the eggs in capsules and taking care of brood until hatching and pigmentation of the larvae. It was concluded that behavioural characters are difficult to interpret in a phylogenetic context due to convergence. However, in some instances they provide valuable clues to reconstruct the evolution of a group and give a good phylogenetic signal.
Morphology
Wing folding mechanisms have been suggested to be a character with phylogenetic value at higher taxonomic ranks (Hammond 1979). Differences in the structure (presence of patches of microtrichia) and mechanism (abdominal movements helping with folding) of wing folding among lineages of Carabidae are not congruent with phylogenetic inferences derived from other characters. The Trachypachidae is a lineage distinct from carabids, a conclusion congruent with recent molecular (Maddison et al. 2009) and karyotypic data (Martínez-Navarro et al. 2011), whereas Gehringia was close to other carabids, as currently accepted. The basis for investigating the phylogenetic value of wing venation within Adephaga and Carabidae was outlined by Ward (1979). This topic has received little attention, perhaps because there is a generalised model in Carabidae that shows a relatively low degree of variation within particular lineages at the tribal or generic level.
Higher-ranked taxa were considered by Beutel (1998) when analysing the relationships of Trachypachidae, based on morphological and functional characters of adults and larvae. He concluded that the family Gyrinidae is sister to all other Adephagan groups. Of these clades, Haliplidae was sister to the remaining families; these were in turn split into two main clades, one made up of Carabidae (including Rhysodini and Cicindelitae), the other made up of (Trachypachidae) + (Noteridae(Amphizoidae+Dytiscidae)). These results contradict Beutel and Haas (1996), who found Trachypachinae to be sister to Carabidae; Beutel and Haas’ hypothesis has recently received support from molecular analyses (Maddison et al. 2009). Ancestors of Adephagan beetles were probably associated with riparian habitats and it has been postulated that independent colonisations of aquatic habitats gave rise to the families Gyrinidae, Haliplidae and Dytiscidae.
Liebherr and Will (1998) studied the phylogenetic value of characters of the female reproductive tract at an inclusive scale that covered the whole family Carabidae. Surprisingly no character defined the Carabidae as a monophyletic taxon; instead the Isochaeta appeared as the adelphotaxon of Anisochaeta (that included Gehringiini and Rhysodini). In turn, the Anisochaeta was divided into two clades separated by the evolution of a secondary spermatheca. Less inclusive clades within these two major groups of Anisochaeta showed relationships that agreed with previous hypotheses in some cases.
Arndt (1998) analysed the phylogenetic relationships derived from larval morphology in 44 tribes of Carabidae. He found support for a monophyletic Carabidae+Tachypachidae+Dytiscidae clade. The family Carabidae was also a monophyletic clade if Rhysodidae were excluded. The Cicindelitae was also monophyletic and showed several autapomorphies. Metriitae and Paussitae made up a monophyletic clade. The subfamily Harpalinae (“higher” carabids) appeared to be a monophyletic clade but relationships of Brachinitae were ambiguous and remain a major challenge for future studies; a close relationship with Harpalinae is unlikely.
The phylogenetic relationships among basal grade Carabidae was revisited by Kavanaugh (1998) who showed that Trachypachidae is sister to all carabid taxa examined (which confirms similar conclusions reported in former works), that the supertribe Nebriitae is a grade rather than a clade (Nebriini is separated from related tribes), and that cicindines are related to Carabini, Cychrini, Cicindelini and Omophronini.
Cladistic analyses based on different data sets (morphology, ethology, geographic distribution), were carried out to investigate the phylogeny of Paussinae (Nagel 1979b), Ozaenini plus Metriini and Paussini (Vigna Taglianti et al. 1998), the Agra cayennensis group (Erwin 1996), the supertribe Nebriitae (Kavanaugh 1996), the subtribe Calleidina (Lebiini; Casale 1998), the Western Hemisphere Pseudomorphini (Erwin and Geraci 2008), the tribe Rhysodini (Bell 1998; which is likely a highly specialised predator of slime moulds rather than a primitive Adephagan stock), the subfamily Broscinae (Roig-Juñent 1998), and the subfamily Psydrinae (Baehr, 1998). These studies either corroborated previous ideas about relationships of taxa or shed light on new and unsuspected hypotheses about the phylogeny and classification of taxa, including the erection of new high-ranked taxa.
Defence substances
Characterisation of chemical compounds used for defence and the phylogenetic interest of this trait was summarised by Moore (1979). The review showed that (i) compounds can be grouped into at least nine categories according to their chemical nature, (ii) there probably occurred a convergent development of the same substances in distantly related lineages, (iii) diversification of chemical types occurred within some subfamilies (e.g. Pterostichinae) whereas others (Harpalinae, Lebiinae) are much more uniform; (iv) the phylogenetic signal of this trait is valuable at tribal level or higher ranks; some compounds seem to vary in particular lineages (Australian Panagaeninae) and could be useful for assessing relationships at lower ranks; and (v) further insight into this trait would result from the study of biochemical synthetic pathways, fine structure of defensive glands and the detection of more subtle compounds.
Karyotypic evolution
A number of contributions have addressed the question on the ancestral karyotype of Adephaga and the Carabidae, and its main patterns of evolutionary change (Nettmann 1986; Serrano 1986; Serrano and Galián 1998), or referred to the karyotypic evolution of particular taxa (Harpalini: Serrano et al. 1994). The family Carabidae (915 taxa analysed) is characterised by a notable variation of the diploid number (2n = 4 - 69), the occurrence of high chromosome numbers in comparison to Polyphagan beetles, and a repeated karyotypic formula in well-studied lineages (e.g. 2n = 26 + XY in Carabini; 2n = 22 + XY in Bembidiini; 2n = 36 + X in Harpalini).
The ancestral karyotype of Coleoptera, still present in many Polyphagan lineages, 2n = 18 + Xyp, had probably undergone significant changes in the ancestors of carabids, since neither this number of autosomes nor the particular Xyp sex chromosomes are found in any carabid. The ancestral condition of a 2n = 36 + X0 male karyotype is widespread in many lineages and may be notably diversified in particular carabid lineages. The occurrence of this formula in some dytiscids and in trachypachids (Martínez-Navarro et al. 2011) provided further support to this hypothesis. However, it has not been found in lineages showing plesiomorphic morphological characters, which suggests that it has evolved rapidly in earlier offshoots of the Carabidae.
Karyotypic data have been shown to be valuable for understanding carabid systematics though it seems that karyotypic changes are not a main driving force for speciation in carabids. This is not to deny the role of karyotypic changes in reinforcing isolation mechanisms in recently originated taxa, regardless of the occurrence of speciation processes under conditions of geographic isolation or in lowland areas (Serrano 1992).
Serrano et al. (1994) summarised the karyotypic data of members of the tribe Harpalini, and found that ancestors likely had a 2n = 36 + X male karyotype. Constraints to numerical variations within this tribe are similar to those found among other carabid tribes. The Ditomina are peculiar because they show high chromosome numbers, which corroborates its ranking as a separate subtribe.
Molecular data
The number of molecular studies have increased since the 1990s, either based only on molecular data or (more recently) combined with other data sets. Inferred relationships have corroborated relationships derived from traditional taxonomy but also often contradicted these, thus emphasising the need of more holistic approaches aimed at obtaining robust and congruent phylogenies.
Maddison et al. (1998) published the first comprehensive DNA-based phylogeny of Carabidae. They studied the nuclear small subunit (18S) ribosomal DNA, sequenced in 35 carabid genera representing 26 tribes. All higher-level clades were monophyletic except for the Scrobifera (scaritines plus clivinines); the Trechitae was sister to Patrobines; Morion and Pseudomorpha were members of Harpalinae; Psydrus and elaphrines were sisters and both were sister to trechites plus patrobines; there was a grade including scaritines immediately below Harpalinae.
A combined analysis of larval morphological characters and molecular data of Cicindelitae showed a number of inferences that contradict current systematics: Omina had a basal position, Megacephalini was a polyphyletic taxon, and Cicindelinae was not monophyletic (Vogler and Barraclough 1998). Use of the resulting inferences showed that there are differential diversification rates among major lineages (e.g. a high rate of diversification was found at the base of megacephalines and collyrines, and another at the base of cicindelines).
Düring and Brückner (2000) investigated the phylogeny and history of lineages of Molopina using molecular analysis based on the sequence of two mitochondrial DNA fragments. Representatives of the genera Percus, Molops and Abax were included, as well as Pterostichus and Carabus as outgroups. These three genera made up a monophyletic clade, and Molops and Abax were sister taxa. In a further step, Brückner and Mossakowski (2006) investigated the phylogeny of the genus Percus by integrating previous molecular, morphological and biogeographic characters with those of nuclear 28S rRNA. This genus is likely a monophyletic taxon divided into three main clades. Relationships among the Tyrrhenian taxa remained unresolved probably as a result of recent diversification and low mutation rates of the molecular marker.
A molecular study of the tribe Harpalini based on the mitochondrial cox1 gene (Martínez-Navarro et al. 2005) showed that (i) Pelmatellina should be included within Stenolophina, (ii) subtribe Harpalina is polyphyletic, (iii) Ditomina is a valid subtribe, and (iv) Selenophori should be ranked as a valid subtribe closely related to the Anisodactylina.
An analysis based on sequences of 28S and wingless genes of Ildobates neboti (a rare hypogean species inhabiting a few caves in eastern Spain) and related taxa showed that tribes currently included in Dryptitae (Dryptini, Galeritini and Zuphiini) made up a monophyletic clade, and that Ildobates neboti is a member of the Zuphiini (Ribera et al. 2006).
Vogt et al. (2005) studied the relationships of African Anthia and Termophilum, and the related Cypholoba chaudoiri, based on the sequence of the mitochondrial nd5 gene. Taxa of Anthia made up a monophyletic clade in which Cypholoba chaudoiri was unexpectedly included. Taxa of Termophilum made up two distinct clusters, which suggests paraphyly of this genus.
Current division of the genus Calathus (Sphodrini) was investigated on molecular grounds by sampling a cox1-cox2 fragment in 44 taxa (Ruiz and Serrano 2006). The monophyly of the subgenus Calathus was corroborated, as well as the distinctness of the monotypic subgenera Bedelinus and Iberocalathus. The subgenus Neocalathus is polyphyletic and needs taxonomic revision and the same holds true for the Canarian Lauricalathus. The latter subgenus should be divided into two subgenera, and one of these should include Trichocalathus.
3 Biology
3.1. Life history strategies and rhythms
Land animals evolve strategies to optimise and synchronise their life cycle with seasonal changes of the environment. For example, reproduction usually takes place under optimal conditions, while metabolism may be reduced if conditions are suboptimal (e.g. dormancy, which in carabids has thus far only been observed for larval and adult stages).
Ultimate (limiting) factors regulating ground beetle life histories
Ultimate factors determining beetle life cycles include variation in temperature and rainfall. Optimal development of the immature stages requires an estimated temperature range of 4–35°C. Rainfall, in combination with temperature, affects soil humidity, which is critical because eggs absorb water from their surroundings to complete embryonic development (Paarmann 1986) and larvae are sensitive to desiccation (Paarmann 1973).
Food can also be critical. Reproduction of, for example, seed-feeding carabid species may be governed by ripe seeds that usually appear at the end of the wet or warm season. Only very few habitats offer suitable conditions for polyvoltine development throughout the year, for example, lake shores, swamps and some lowland rainforests with very short dry spells.
The only ultimate factor determining carabid beetle life cycles in the Arctic, Subarctic and Antarctic, as well as in montane habitats of the temperate zone is temperature (e.g. Thiele 1977). In the summer, only a short time window exists for reproduction and development. All species in these habitats are summer developers. Species with rapid larval development, such as Pterostichus adstrictus (Paarmann 1994), are true summer breeders with adult hibernation only. Species with slow larval development hibernate as larvae as well as adults and require more than one season to complete their life cycle (Kaufmann 1971; Davies 1972; Matalin 2008). In the mountains of temperate Europe (altitude of 2200–2600 m) the favourable season is reduced to 3–4 months. In forest Pterostichines, especially in the large genus Pterostichus and in the Molopines Abax and Percus, cycles are often biennial (Brandmayr 1977). In the genus Molops, where embryonic development can last for more than one month and the eggs are guarded in a subterranean hole, the females disappear from the soil surface during summer, and reappear in the autumn. The subterranean larvae are active during winter, and the new generation requires a further year to reach maturity (Brandmayr and Zetto Brandmayr 1991).
Larsson (1939) recognised different reproductive strategies in carabid beetles of the temperate zone by studying museum material. He divided them into Frühlingstiere (spring breeders) and Herbsttiere (autumn breeders). Spring breeders reproduce during the spring and hibernate as adults only. Autumn breeders reproduce during the autumn and hibernate mainly as larvae. In a number of species, adults may hibernate after reproduction to enter a second reproductive period (Gilbert 1956; Vlijm et al. 1968; Schjøtz-Christensen 1968; Krehan 1970). (Schjøtz-Christensen (1965, 1966) showed that in some Harpalus species spring and autumn breeding populations co-occure in the same habitat. Other examples include Abax parallelepipedus (Löser 1970), Poecilus lepidus (Paarmann 1990), Pseudophonus rufipes (Matalin 1997a) and Harpalus affinis (Matalin 1998). A third breeding category – spring-autumn breeder (Matalin 1997b) – is found in the genus Stenolophus. In 1990 Den Boer and Den Boer-Daanje, summarising the life history strategies of 68 common carabid beetles in Drenthe (the Netherlands), found a continuum of reproduction from early spring to late autumn, and seven of them reproduced during winter. Den Boer and Den Boer-Daanje distinguished species with summer larvae (summer developers, 40 species) and species with winter larvae (winter developers, 28 species). Drenthe is located in an area with Atlantic climate: warm winters and wet summers, thus offering a broad reproductive window. In areas with a continental climate, however, this window is much narrower.
Cave environments are buffered against climatic variation and can have (i) a constant temperature throughout the year, or (ii) distinct seasonality. Trechines living in caves are mostly autumnal reproducers with winter larvae. The rhythms of Aphaenops and related genera may show distinct seasonality at least in the activity of adults, influenced by the cave’s air humidity (Juberthie 1969), and sometimes with two distinct annual peaks (Cabidoche 1963, 1966). Reproduction may coincide with a peak in food, as found between Neaphenops tellkampfi and the eggs of the orthopteran Hadenoecus subterraneus (Kane et al. 1975).
The seven winter breeding species found in the Netherlands (see above) connect the carabid fauna of the temperate zone with the life history strategy typical for the subtropics with winter rain. In Palestine, Bodenheimer (1934) only caught beetles from October to June. Winter breeding (rainy season breeding) is a typical reproductive strategy in habitats that are dry in the summer, such as North Africa (Paarmann 1970, 1975). In specific habitats with moist soil during the dry summer period, propagation and reproduction occur throughout the year (Paarmann 1975, 1976d). Thermophilum sexmaculatum and Graphipterus serrator, with specialised larvae that feed on ants and their brood, reproduce in the summer (Paarmann 1985; Paarmann et al. 1986; Dinter et al. 2002), but only in sandy soil that acts as a moisture trap.
In Mediterranean Europe, which is also dry in the summer, some seed-feeding carabids - the ditomines Carterus calydonius, Ditomus clypeatus, and harpaline carabid beetles that provide Daucus or Plantago seeds to their larvae (Brandmayr and Zetto Brandmayr 1974; Schremmer 1960) - show summer reproduction. Other seed-feeders (Ophonus, Pseudoophonus) are adapted to more humid soils and normally reproduce in the autumn (winter larvae; Zetto Brandmayr 1983a, b).
No information is available on the reproductive strategies of Carabinae from the subtropics with summer rain. However, it seems reasonable to suggest that they show rainy season propagation (summer breeding) in habitats which are dry during winter. A number of studies on carabid beetle life histories are available from the tropics. In Central Africa (Kivu district), which is characterised by low variation in median air temperature (0.9 °C) and low rainfall from June-August (Walter and Lieth 1960), the majority of species avoid reproduction during and around the dry season (Paarmann 1976b). Dry season propagation was only found in two species, one living in a swamp and one in a cultivated area. North Sulawesi (Indonesia) is without a dry period, yet the appearance of gonad dormancies was widespread among 155 carabid beetle species: 65% had at least one dormant female (Paarmann and Stork 1987; Stork and Paarmann 1992). Females of the canopy dweller Colpodes buchanani also synchronise reproduction with annual temperature changes typical of the subtropical climate (Paarmann and Paarmann 1997).
Along the Amazon River in Brazil, forests are often inundated for up to seven months of the year. This flooding is independent of the rainy season in central Amazonia. During flooding, carabid beetles occur on tree trunks or in the canopy in the inundated site, reproducing when the water level is low (Adis et al. 1986; Adis et al. 1990). In lowland rainforests, carabids aggregate in areas with an accumulated amount of organic matter, such as fruit falls (Erwin 1979b). These fruit falls are unpredictable in space and time, lasting only for a few weeks. Fig fruit falls play an important role in these rainforests, as they occur virtually throughout the year. Distinct carabid assemblages have been found at fig fruit falls in lowland rainforests of the Amazon basin (Paarmann et al. 2001), Brunei (Borcherding et al. 2000), Australia and Africa (Paarmann et al. 2006). Female gonad maturation starts immediately after locating a fruit fall, with some females carrying ripe eggs combined with the undeveloped ovaries. These ‘transport eggs’ can be deposited directly after arrival at the fruit fall, providing larvae more time for development. While moving between patches of fruit fall, females experience short gonad dormancy induced by food shortages (Paarmann et al. 2001; Arndt and Kirmse 2002).
Proximate factors and endogenous rhythms
During unstable temperatures, soil humidity and resources, proximate factors and endogenous rhythms play a major role in controlling carabid beetle life cycles. At temperate latitudes, many species, especially species with summer larvae, use photoperiodic changes to synchronise gonad maturation (Thiele 1977). Autumn breeding species display thermic parapause (Müller 1970): an obligatory dormancy at a genetically fixed developmental stage, where the phase of induction cannot be recognised. Larval development can only be completed after passing a certain period of time at low temperatures. Larvae of other species with winter larvae, such as Abax ovalis and Abax parallelepipedus, only pass a thermic quiescence (Müller 1970): a facultative delay or suspension of development. This may also be the case for species with winter larvae at higher latitudes (and montane regions): Subarctic populations of Pterostichus nigrita were still under photoperiodic control in terms of gonad maturation, yet displayed a shift of the response curve to longer day lengths (Ferenz 1975).
Annual day length amplitudes decrease from higher latitudes to the equator, as does the importance of photoperiodic changes as a proximate factor. However, day length changes of 1 h can control imaginal diapause (Norris 1959, 1965). Two carabid species from North Africa synchronise their life cycle with annual rainfall, triggered by a decrease in temperature and a decline in the photoperiod (Paarmann 1974, 1976c). This control mechanism in a rainy season breeder (or winter breeder) of the subtropics with winter rain shows marked similarities with temperate autumn breeders and aestivation (Thiele 1977).
In the Kivu region, Central Africa (see Paarmann 1976b), the maximum change in daylight is 16 min only, and the maximum annual temperature change is 0.9°C. Under such climatic conditions, temperature plays a role as a proximate factor. The temperature of the upper soil layers and the soil surface is influenced by the water content of the soil. With water loss in the upper soil layers, daily temperature fluctuations increase. Some hours of higher temperatures per day induce gonad dormancy. With the onset of rainfall, temperature fluctuations decline and dormancy is terminated. Synchronised maturation is stimulated by the increase in average temperatures (Paarmann 1986).
Endogenous control of gonad dormancies
The synchronisation of gonad maturation with seasonal change in ultimate factors is possible only if proximate factors influence the endocrine system controlling this maturation. Emmerich and Thiele (1969) and Hoffmann (1969) were the first to study the hormonal control of gonad maturation in spring breeders. They found a connection between proximate factors, neurosecretions and the activity of the corpora allata, which produces juvenile hormones (JH). JHs are necessary to complete gonad maturation in males (Ferenz and Hölters 1975). In females, only previtellogenesis is controlled by JHs. To complete ovarian maturation, the production of a second hormone is postulated. Applications of JHs to dormant beetles of the winter breeder Orthomus barbarus have confirmed a similar control mechanism for this breeding type (Paarmann 1976a). The same application to dormant beetles of the summer breeder Pogonus chalceus resulted in complete maturation of both sexes, even complete gonad maturation in females, meaning that either complete maturation is controlled by JHs only, or high temperatures suppress only the production of JHs but not of vitellogenic hormones.
Endogenous rhythms are involved in gonad maturation. Under constant environmental conditions gonad maturation is controlled by an endogenous rhythm, synchronised by an external cue such as soil temperature (Paarmann 1986). In the desert-dwelling carabid beetle Thermophilum sexmaculatum thermoregulational behaviour is controlled by a circannual rhythm, resulting in lower body temperatures at the end of the optimal reproductive period, which causes an inactive stage of the gonads (Erbeling and Paarmann 1986).
As part of the taxon pulse theory (Erwin 1979b), ground beetles from tropical areas undergo latitudinal and altitudinal expansion, leading to climatic specialisation, including the development of dormancy to survive unfavourable climatic conditions. If all carabid beetle dormancies are based on a uniform hormonal system, manifold convergent evolution is possible. The use of gonad dormancies to synchronise life cycles with changing environmental conditions is widespread among tropical carabid beetles. Only one Abacetus species, living under stable humidity and temperature conditions (the shore of Lake Kivu, Central Africa), seems to develop without dormancy. With the exception of short gonad dormancies, triggered by food shortages in the seed-feeding guild, all studied gonad dormancies are under the control of temperature as a proximate factor.
Specialisation along riparian habitats (pathway i) leads to a synchronisation of the life cycle with seasons with stable moisture conditions, especially along riverbanks. Specialisation in seasonally dry habitats (pathway ii) leads to a synchronisation of the life cycle with the period of optimal soil humidity, e.g. rainy season propagation (Paarmann 1979). While larvae of winter breeders in the subtropics with winter rainfall are adapted to comparable temperatures, a small group requires high temperatures for successful development. These specialists, whose larvae feed on ants and ant brood, have evolved along pathway (ii) in the subtropics with summer rainfall and spending the winter in gonad dormancy. Such species have yet to be reported in the temperate zone.
Larsson (1939) found no autumn breeders among 21 studied species of the old genus Agonum. These species are possibly all descendants of one common ancestor that reached the temperate zone along pathway (i) after which some descendant species adapted to non-riparian habitats. One member of this group, namely Platynus (Agonum, Limodromus) assimilis is a spring breeder, but its gonad dormancy is controlled in a fundamentally different way than in other spring breeders, by a photoperiodic quiescence (Neudecker and Thiele 1974).
Gaps in our current understanding of carabid beetle life history strategies include (i) a lack of knowledge on life history strategies in the subtropics with summer rainfall, in the tropics with long dry seasons and in areas with unpredictable rainfall, (ii) whether canopy dwelling carabid beetles in tropical rainforests display seasonal patterns, and (iii) a detailed study on the hormonal control of dormancies in carabid beetles, as no such studies have been performed since Ferenz (1977).
3.2. Carabid beetle food
Carabid beetles are generally considered polyphagous predators. However, in line with their enormous species richness and diversity in body shapes and biotopes they inhabit, a whole range of trophic specialisations occurs in the Carabidae (Hengeveld 1980a; Zetto Brandmayr et al. 1998b). Although carabid feeding ecology and biology has been studied frequently (also during ECM meetings), it is surprising how many basic questions on carabid food remain unanswered. Except for Larochelle (1990), who mentioned food preferences of 1054, mainly North-American, European and Japanese species, basic information on food preferences or requirements is often lacking, even for many common species. This chapter does not attempt to review all trophic specialisations of Carabidae; it has been done before (Thiele 1977; Hengeveld 1980a; Toft and Bilde 2002). Instead, it focuses on recent advances in the domains of seed and ant feeding, as well as unique life history strategies, such as ectoparasitism and the predation of amphibians.
Seed feeding
Carabid beetles accept a variety of plant foods such as leaves, fruits, pollen, seeds and fungi (Toft and Bilde 2002 and references therein). Seed feeding, or granivory, occurs in many species including polyphagous ones that prefer animal prey (Lund and Turpin 1977; Hengeveld 1980b; Toft and Bilde 2002). True granivory, i.e. where seeds are central to the species’ food budget, has evolved in two tribes of Carabidae, Zabrini and Harpalini. The ecology of granivorous carabids is of great interest since granivory required the evolution of morphological, physiological and behavioural adaptations associated with crushing, digesting and foraging for seeds. To crush hard seeds, adults and larvae of granivorous species have evolved broad mandibles with massive adductors (Zetto Brandmayr et al. 1998b; Paarmann et al. 2006). Sclerotised structures in the adult proventriculus are then used for fine grinding of the ingested seed fragments (Evans and Forsythe 1985). Behavioural adaptations have involved, for example, climbing plants and storing seeds in burrows (Thiele 1977). Physiological adaptations to seed feeding are understudied but recent evidence shows that digestion of seeds is facilitated by endosymbionts (Lundgren and Lehman 2010).
The amount of seeds eaten by carabids in the field may be substantial. Based on seed losses of artificially exposed seeds, Honek et al. (2003) estimated that up to 4000 seeds m-2 d-1 may be removed by carabid beetles in arable fields in the Czech Republic. Honek et al. (2005) reported that carabids, mainly Amara montivaga, destroyed about 83–88% of the annual seed production of Taraxacum officinale spp. agg., and Kjellsson (1985) showed that approximately 65% of the annual seed production of Carex pilulifera L. was consumed by a single species, Harpalus solitaris. However, individual capacity for eating seeds varies with season (Honek et al. 2006) as a result of natural phenological changes (transition from dormancy to reproduction, dispersal, breeding and searching for overwintering sites). Consumption is also affected by temperature (Saska et al. 2010). Clearly, carabid beetles may have an important impact on the reproductive success and dispersal of plant species, but more research is needed on how these affect the population dynamics of plants in the longer term. Larvae should also be considered in these studies, as their consumption of seeds can be comparable to that of adults (Klimeš and Saska 2010).
The consumption of particular seed species is ultimately determined by the preferences of the carabids in question. During the last 30 years, a number of authors have investigated carabid preferences for seeds in the laboratory using choice (cafeteria) experiments (Lund and Turpin 1977; Brust and House 1988; Jørgensen and Toft 1997a). Most studies, however, have established preferences based on a limited number of seed species (usually 2–5). Only Honek et al. (2003, 2006, 2007) tested seed preferences in carabids using 64 or 28 species of herbaceous seed. Honek et al. (2003, 2007, 2011) demonstrated that the preference for seeds correlates with carabid body size: on average, smaller species prefer smaller seeds, and vice versa. Larger carabids also consume a greater variety of seed species and Harpalini are less specialised than Zabrini (Honek et al. 2007). However, there are other characters such as seed shape, thickness of the testa (Lundgren and Rosentrater 2007) and nutrient content of the seed that affect preference. Similarly to other seed-cracking organisms (e.g. Diaz 1994), mandible size and shape determine the seed preferences of Notiobia species occupying fruit fall sites in tropical forests (Arndt and Kirmse 2002; Paarmann et al. 2006), and these preferences are consistent throughout the season (Honek et al. 2006).
Taxonomic affiliation constrains the preferences for food in many insect groups. Earlier research as well as direct field observations have indicated that species of certain genera had specific affinities with respect to their seed preferences. For example, Brandmayr and Zetto Brandmayr (1987) and Zetto Brandmayr (1990) suggested that most Ditomina and Ophonus (both Harpalini) are associated with Apiaceae, while Harpalus (Harpalini) is unspecialised in this sense (Zetto Brandmayr 1990). Hurka (1996) reported that species of the subgenus Zezea (Zabrini: Amara) may be associated with Poaceae. The existence of a taxonomic constraint has been experimentally confirmed by Honek et al. (2007), who carried out a cafeteria experiment that included 28 seed species and 30 carabid species. They demonstrated that species of Zabrini mostly prefer seeds of Taraxacum, while species of Harpalini prefer seeds of Cirsium and Viola. Carabids not only distinguished seeds from different families, but they were also able to discriminate between seeds at a finer taxonomic scale, i.e. seeds of different sections of the Taraxacum officinale species complex (Honek et al. 2011). The origin of seeds plays a role in some carabid species. For example, Honek et al. (2011) fed Czech carabids with Italian and Czech seeds of the same plant species and found that the beetles preferred the latter. It is likely that the existence of specialisation on particular seeds reduces the competition for food and allows the coexistence of species in the same habitat.
Seeds are nutritious, but their value as food for carabids has not been appropriately recognised until recently. The value of food is best defined by its contribution to the fitness of the consumer (Toft and Bilde 2002). Fitness parameters that are commonly used as criteria for the evaluation of food quality are female fecundity, survival and duration of larval development, and the attainable body size. Zetto Brandmayr (1976) showed better survival in larvae of several species of the genus Ophonus when provided with seeds of Apiaceae compared to other seeds or insects. Although Jørgensen and Toft (1997a, b) stimulated further research on this topic (mainly in Europe and Japan), information on how seed diet affects fitness is only available for a small number of species. Adaptations to granivory have evolved to varying degrees in different taxa, and even closely related species may show different strategies (for Amara, subgenus Amara, compare e.g. Jørgensen and Toft 1997b; Saska and Jarošík 2001; Hurka and Jarošík 2003; Fawki and Toft 2005; Saska 2008; for Amara, subgenus Curtonotus, compare e.g. Saska 2005; Sasakawa 2007; 2009); for Notiobia, see Arndt et al. 1996; Paarmann et al. 2001; Arndt and Kirmse 2002). More interestingly, particular seed diets may have contrasting effects on different fitness traits (Fawki and Toft 2005). The effects of maternal diet (Saskawa 2009) or diet of the previous generations (Hurka and Jarošík 2003) on larval performance are poorly studied. Also, worthy of mention here is the scoring system of Paarmann (2002) used to evaluate larval performance under different dietary regimes. In general, larvae are more specific in their food preferences than adults (Thiele 1977) because of increased selection pressures on larvae (Sasakawa 2007) and due to morphological constraints on the suitability of the available food during the early stages of development (Paarmann et al. 2006). Klimeš and Saska (2010) argued that this selection pressure is highest in the first instar larva and decreases in older instars, with increasing the head width/seed size ratio in larvae and widening the range of edible food items.
Ant feeding
Ants are the most abundant group of organisms on Earth in terms of biomass (Hölldobler and Wilson 1990). Not surprisingly they represent an important food source for many other taxa, including carabid beetles. Polyphagous carabid species frequently prey on ants (Thiele 1977; Hengeveld 1980b), and several clades have adapted to ant feeding with some having evolved the highest degree of specialisation, i.e. myrmecophily. In general, biological information is very limited and needs systematic study.
Species that have adapted to feeding on ants have evolved interesting behavioural and morphological adaptations, including chemical mimicry that reduces the risk of being attacked by their hosts (Zetto Brandmayr et al. 2000a; Dinter et al. 2002). Larvae of Sphallomorpha (Pseudomorphini) form burrows close to ant nests and attack ants that pass by (Moore 1974). Associations with ants and termites seem to be a joint character for the entire tribe of Pseudomorphini, though evidence is limited (Baehr 1994). Species of the Siagonini also prey on ants ( 1998a, ). Species of the genus Siagona inhabit crevices in the soil near ant nests and attack ants both as adults and larvae, but do not seem to enter ant nests frequently (Bauer et al. 2005). The larvae of some Ozaeini use so-called terminal disks (modified last abdominal segments) for attracting and capturing ants (Di Giulio and Vigna Taglianti 2001; Moore and Di Giulio 2006).
Adults of the North African Anthiini and Graphipterini are free-living but larvae enter ant nests where they prey upon ants to complete their development (Paarmann 1985; Paarmann et al. 1986). The larva of Thermophilum (Anthiini) moves freely in the nest after it gains chemical mimicry from ants it has previously attacked (Dinter et al. 2002), and consumes both ants and ant brood (Paarmann and Erbeling 1986). In contrast, the larva of Graphipterus serrator forms a chamber inside the ant nest where it stores ant brood before consumption, and hides against ant attacks (Dinter et al. 2002). Species of Thermophilum, as well as Graphipterus serrator, show preferences for particular ant species, Graphipterus being the least selective (Dinter et al. 2002).
True myrmecophily (and perhaps termitophily) evolved in the tribe Paussini, in which morphological and behavioural adaptations are prominent in both adults and larvae (Nagel 1979b; Di Giulio and Moore 2004; Moore and Di Giulio 2006). Although this association is well known, data on food requirements or trophic associations are known for a limited number of taxa only, and this requires further investigation.
Unique life history strategies – ectoparasitism and the predation of amphibians
The variety of life history strategies in carabid beetles includes ectoparasitoidism, a strategy otherwise rare in beetles. Parasitoids are insects whose larvae develop at the expense of a single prey individual (a host), which ultimately dies as a result of parasitoid feeding (Vinson 1976). Ectoparasitoid larvae attach to the host body and feed externally on it, while their adults are free-living (Vinson 1976).
Ectoparasitoidism has been described from four carabid genera: Brachinus (Brachinini), Pelecium (Peleciini), Lebia and Lebistina (both Lebiini) (Weber et al. 2008), but several related genera show tendencies towards parasitoidism (Erwin 1979a; Frank et al. 2009). The life cycle of a typical carabid ectoparasitoid includes (i) a female depositing eggs in the host habitat when hosts are present; (ii) mobile early instar larva searching for and attaching to a suitable host; (iii) after attachment, a short physogastric feeding phase, typically with rapid ingestion; and (iv) a distinct pre-pupal “resting” phase during which the host is consumed.
Despite the early discovery of ectoparasitoidism in Carabidae (e.g. Wickham 1893; Silvestri 1904), known host associations are few. With one known exception, beetle pupae are the hosts. Larvae of Lebia (five species known to be parasitoids) and Lebistina (one species) parasitise leaf beetle (Chrysomelidae) pupae (Weber et al. 2008). Larvae of a single undetermined species of Pelecium have been observed developing on chrysomelid pupae and millipedes (Salt 1928). Nearctic wetland species of Brachinus (seven species) parasitise the pupae of water beetles (Dytiscidae, Gyrinidae, Hydrophilidae) (Saska and Honek 2004). Despite suggestions proposed by Jeannel (1942), the discovery of the hosts for dryland species of Brachinus from Europe was only made 60 years later. Saska and Honek( (2004, 2005) successfully reared two species (Brachinus explodens and Brachinus crepitans) on the pupae of another carabid genus, Amara, a finding that has recently been confirmed for Brachinus elegans by Makarov and Bokhovko (2005).
Besides direct observations, host-parasite associations have frequently been suggested simply on the basis of co-occurrence of the carabid parasitoid and potential host species. In some cases, however, these observations have led to erroneous predictions (Jeannel 1942; Perez-Zaballos 1985), subsequently refuted because the life cycles of the two suggested partners are not synchronous. Such synchrony has so far been demonstrated only for Brachinus explodens and Brachinus crepitans (Saska and Honek 2008). Thus, when looking for hosts of Mastax or Aptinus (both Brachinini) or wetland Palaearctic Brachinus species, both co-occurrence and synchrony should be taken into account. More discoveries are probably to be made in the tropics, as that climatic zone contains a vast diversity of lebiine carabids (Ober and Maddison 2008). Research is also needed on the ecology of ectoparasitic carabids to determine the adaptive significance of life history traits of this peculiar strategy. In most cases, available information relates to a brief description of development; only a few species have been studied in detail (Erwin 1967; Juliano 1985; Saska and Honek 2004, 2005, 2008; Weber et al. 2006). Host selection, food utilisation or the adaptive significance of variation in the number of instars (2–5 instead of the typical 3) could produce interesting results. Mimetic complexes have been described between adults of Lebia and chrysomelids, including species for which parasitoidism is unknown (Hemenway and Whitcomb 1967), suggesting further trophic associations between the two groups. Focusing on taxa representing transitional evolutionary steps to parasitoidism (Erwin 1979a; Frank et al. 2009) may shed light on the evolution of parasitoidism in Carabidae and in Coleoptera in general.
Carabid beetle larval and adult predation on amphibians has recently been described in Israel. Elron et al. (2007) have shown that larvae of the carabid Epomis dejeani preyed upon two amphibian species (Bufo viridis and Hyla savignyi), confirming an earlier brief note by Moore (1971) from Australia. Subsequently, Wizen and Gasith (2011) performed laboratory experiments, and showed that adults of the two sympatric Epomis species in Israel, Epomis dejeani and Epomis circumscriptus, prey upon five and four amphibian species, respectively. Wizen and Gasith (2011) argue that little is known about the feeding habits of sympatric congeneric insects, and that the partial food overlap of these Epomis species warrants further investigation.
3.3. Dispersal
Carabid beetles found a rich niche in ecological research through the peculiarities of their dispersal power. Sahlberg (1868) recognised that carabid species exhibit a variety of wing attributes, including wing dimorphism, and that this has implications for their powers of dispersal. Darwin was probably the first to consider the evolutionary and ecological implications of wing polymorphism in Coleoptera after recording high proportions of flightless beetles on the island of Madeira. He hypothesized that flight ability might be evolutionarily disadvantageous for species from insular populations, as they would be more likely to get carried away from the island (Darwin 1859). A few decades later, Darlington turned his attention to the low proportions of macropterous carabids in isolated locations such as islands and mountain tops, and concluded that wing reduction must confer enhanced viability (Darlington 1936, 1943). Lindroth (1988, 1992a, b) studied the wing morphology of carabid assemblages from islands in the Baltic Sea in comparison to control assemblages from nearby mainland sites. He found that the proportions of brachypterous and macropterous species were both lower in insular than in mainland assemblages, even whilst macropterous species were predominant in all of the studied assemblages (see also Ås 1984, Kotze et al. 2000). Dimorphic species, on the other hand, were more numerous in insular than in mainland faunas (Lindroth 1988, 1992a, b). These observations were of fundamental importance to Lindroth’s epic zoogeographical studies, published posthumously in 1992 (Lindroth 1988, 1992a, b). After determining the frequencies of the different wing morphologies in populations of wing-dimorphic carabid species across the Fennoscandian region, Lindroth was able to estimate the relative ages of these populations. On that basis, he theorised about the routes of post-glacial colonisation of Fennoscandia by different species. He was subsequently able to divide the fauna into three elements: Wűrm hibernators, immigrants from a southern route to the west of the Baltic Sea and immigrants from the east. Both Lindroth (1988, 1992a, b) and Den Boer (1970) came to the conclusion that macropterous specimens dominate in recently established populations of dimorphic species, which gradually shift to an increasing proportion of brachypterous individuals as these populations grow older. Observations of pioneering populations of the invasive species Pterostichus melanarius in Canada support this model (Niemelä and Spence 1991).
Largely thanks to the work of Piet den Boer and colleagues, subsequent to the Dutch land reclamation projects of the late 1950s, research interest in the dispersal of carabid beetles flourished, and this provided the theme for the first meeting of European carabidologists in Wijster in 1969, which Piet den Boer hosted (see above). Dispersal power was also the theme of the subsequently published proceedings volume, edited by Den Boer (1971, see also Table 2).
Lindroth was keen to determine the genetic mechanism behind wing dimorphism and conducted breeding experiments with the wing-dimorphic species Pterostichus anthracinus (Lindroth 1988, 1992a, b). The results he obtained, supported by similar results from studies of other coleopteran taxa, led him to conclude that wing dimorphism is inherited in a simple Mendelian pattern, in which brachyptery is dominant. The late Konjev Desender, in whose honour the 14th ECM was held, performed similar breeding experiments using the wing polymorphic species Pogonus chalceus. In this species, crosses between macropterous and brachypterous adults produced offspring with intermediate wing length, suggesting that the genetic control of wing length in this species is polygenic (Desender 1989a). Desender also conducted an exhaustive biometric study of wing development in 300 carabid species indigenous to Belgium and demonstrated that, in addition to brachypterous individuals, also a large proportion of macropterous individuals do not possess functioning flight muscles and are therefore incapable of flight. In the wing-polymorphic Pterostichus vernalis, for instance, some populations are entirely macropterous, with functional flight (but see below) muscles even in relatively short-winged individuals, whereas in some other populations even macropterous individuals lack functional flight muscles (Desender 1989b, see also Nelemans 1987). Desender also studied wing morphology in the genus Calosoma after research trips to Easter Island and the Galapagos archipelago. Three endemic species appeared to be brachypterous, whereas the supposedly introduced species, Calosoma granatense, appeared to be wing polymorphic (Desender et al. 2000).
Berend Aukema conducted breeding experiments with the Calathus melanocephalus group to shed further light on the inheritance of dispersal characteristics. Aukema (1990) demonstrated that these species show a simple Mendelian pattern of inheritance of wing morphology, as described by Lindroth, i.e. simple inheritance with brachyptery dominant over macroptery for the two wing dimorphic species Calathus cinctus and Calathus melanocephalus. However, he also demonstrated that certain environmental factors, such as temperature and food supply, influence expression, with higher temperatures and better food availability resulting in both greater proportion of macropterous individuals (Aukema 1990), and the development of flight muscles (Nelemans 1987). Moreover, long-winged females of these two species had greater fecundity than short-winged females, both in terms of quantity of egg production and duration of egg production (Aukema 1991). This result was somewhat counterintuitive, as a number of other studies of wing dimorphic insects, e.g. Roff (1986) found that brachypterous females are generally more fecund, suggesting that the advantage conferred by brachyptery is enhanced fecundity for females. Furthermore, macropterous females of Pogonus chalceus have greater fecundity, suggesting that long wings and functional flight muscles are associated with large body size (Desender 1989b; Aukema 1991).
Work from other invertebrate taxa has suggested that there is a cost in terms of reproductive capacity for flight, with some macropterous females lysing their flight muscles and shedding their wings prior to reproduction, resulting in enhanced reproductive capacity. Among carabids, Amara plebeja autolyses its wings and can subsequently regenerate them to facilitate migration between breeding and over-wintering habitats (van Huizen 1977, 1979). This is supported by Matalin’s (1994) observation that reproductive females from window traps invariably have fewer ova than those from pitfall traps. Matalin (1994) also concluded that the choice between flying and walking varies considerably between species and with different stages in the life cycle, with flight activity being favoured by dispersive young adults, shortly after emergence and, in Harpalus rufipes and Harpalus calceatus, by mature males. Mature adults exhibit the highest walking activity during the breeding season, apparently being the favoured form of locomotion when seeking a mate (Matalin 1994).
Wing morphology alone is not sufficient to describe dispersal ability in carabids. Desender (2000) and Matalin (2003) studied the phenology of carabids in relation to flight muscle development. Desender (2000) investigated the trade-off between dispersal and reproduction in female carabids from the Belgian fauna, and most of the species he studied supported the oogenesis-flight syndrome, i.e. females with ripe ovaries tend not to possess functional flight musculature. This phenomenon was most pronounced for species that reproduce in late summer or autumn and emerge in late spring (Desender 2000). Matalin (2003) concluded that in females of large species, wing muscles decline during a period of increasing body mass, after development of the gonads.
In addition to the wealth of material on dispersal by flight, carabidologists have also investigated running activity, demonstrating that larger Carabus species run slower than smaller carabids, though in Pterostichinae and Harpalinae, larger species are faster (Mossakowski and Stier 1983). Temperature has a significant effect on running activity in Carabus auronitens (Althoff et al. 1994). Clearly the expression of dispersal ability in carabid beetles is highly complex, being governed by environmental and life cycle factors, in addition to genetic control. It is equally clear that there are still many unresolved issues regarding the dispersal of carabids and we are likely to see studies on this topic at future ECMs. In particular, ongoing land-use change and habitat fragmentation, exacerbated by the influence of climate change, mean stronger selective advantages for species with better powers of dispersal. A major challenge for the scientific community will be to discern evolutionary changes in response to this selective pressure. In conservation, the main challenge will be to develop strategies for the conservation of species with poor powers of dispersal.
4 Methods
4.1. Methodological approaches
Methods influence the way we approach, perceive, and understand the world. All methods have strengths and weaknesses, which make certain things to be easily noticed while others remain hidden or un-emphasised – and such effects of the methods on knowledge often go unnoticed or are unappreciated by researchers. Carabid research has long been dominated by observation and description, but there still remains much to be observed and described about carabids. However, the prevalence of certain methods in carabid research (e.g. pitfall trapping as a collection method, see below) has put a strong stamp on the amount and structure of our knowledge about carabids. Some of the resulting biases are mentioned below; this list is illustrative, not exhaustive.
Prevalence of knowledge about adults: Due to the epigaeic activity of the adults, and the fact that they are more easily collected, manipulated and kept in the laboratory, there is an overwhelming disparity about our knowledge on the ecology of the different life stages of carabid beetles. Our knowledge on carabids was (Lövei and Sunderland 1996) and remains primarily determined by knowledge about adults. A search on Web of Science with the term “carabid* OR ground beetl*” between 2000–2009 yielded 3186 papers, only 460 remaining when this was combined with the term “larv*” (search made by G Lövei, on 4 February 2011).
Geographical unevenness in the origin of our knowledge: This is a general phenomenon: we know that the tropics is more species rich, in general, than the temperate region (already mentioned by Darwin 1859), yet most of our research effort is still directed towards temperate ground beetles. Of the above computer search on ground beetles, only 80 of the original 3186 papers remained when the additional term “tropic*” was introduced. We can safely predict important new understanding emerging from more detailed studies performed in more southerly regions; many of the techniques formerly restricted to developed countries can now be usefully employed in more tropical areas.
Biased perception of carabids as predators: Predators and predation keep us fascinated, possibly because early humans have been both hunters and hunted. However, this colours our perception of the world (see Carabid beetle food above). In the case of carabids, the fact that many species will attack prey offered to them, especially in the laboratory, and that many beetles are indeed fast-moving predators, has led to a widely-held belief that carabids are predators. Carabidologists (mostly) know better, but we have been a bit lax to actively dispel this notion among ecologists, natural historians, and the general public. In relatively recent literature, one still comes across this perception (Braun et al. 2004), and in some cases, elaborate theories are built on such shaky grounds (Lövei and Magura 2006).
The rarity of testable hypotheses: Due to a history of descriptive studies, there seems to be a general rarity of precisely formulated, testable hypotheses. Many studies have the only justification that “we do not yet know, so let’s find out”. With increasingly fierce competition for funding and publication, such arguments do not carry much weight. An additional advantage of formulating hypotheses is that it forces us to think ahead: what is to be expected? Why? However, hypotheses should be well formulated (see Ford 2009; Underwood 2009). In the literature (not only in carabidology) one often encounters the “null hypothesis” formulated as “we expect no differences will be found”. Do researchers really expect that “nothing will happen”? If so, why is the experiment worth performing? Indeed, in the real world the null hypothesis is rarely if ever true as there will always be differences between effects. What is of importance is the magnitude, i.e. effect size, and precision, i.e. confidence interval of the effect (Nakagawa and Cuthill 2007; Läärä 2009). The careful separation of hypothesis formulation vs. the Popperian way of arriving at scientific evidence should not be confused – but often is.
The overall task is unchanged: to understand what made carabids such an evolutionarily successful group. In order to answer this question, one has to quantitatively continue to document the patterns of occurrence of members of this group – this is a logistical, not a methodological challenge. Among the promising “methodological approaches”, modern population genetical toolkits are well used, with several interesting results – it would be good to take these and use them in extra-European habitats as well. Gene expression study methods have recently developed and simplified considerably (Ouborg and Vriezen 2007), and facilitate the study of some interesting ecological questions, such as reaction to such factors as stress and food selection. Modern methods, such as those of ecological immunity, also allow a more refined characterisation of ground beetle reactions to habitat quality.
4.2. Analysing pitfall-trapped carabid data
Pitfall trapping is the best-known collection method used by carabidologists, especially in ecological studies (Lövei and Sunderland 1996). The method, originally described nearly 80 years ago (Barber 1931) and later often referred to as Barber traps (Thiele 1977), is cheap, easy to use and once set up, operates by itself. It allows for adequate replication in field-based studies, and collects large samples (see Fig. 2 for examples of a few commonly used pitfall traps).
Figure 2.
Different pitfall types. A = Jar or yoghurt can. B and C = traps with an outer can to make collecting of the sample easier. B = funnel trap with small jar. C = trap for moist biotopes (the outer can contains gravel or stones to prevent the can from being pushed up by groundwater). V = preservative (usually formaldehyde 3–4% or propylene glycol), S = stones or gravel.
One of the most convenient features of pitfall trapping is also its main disadvantage, because the resulting catch, although beguilingly countable, is not a measure of density, but of activity density. Carabidologists have recognised this and other drawbacks of pitfall trapping, which have often been discussed in the literature from Greenslade (1964), Thiele (1977) and Lövei and Sunderland (1996) to Holland (2002) and regularly at ECMs. However, the method has not been subject to rigorous, thorough testing, nor to a systematic review, and consequently, most carabidologists tip their hat at the problem, then proceed to ignore it, and often use sophisticated evaluation methods to answer important research questions. Needless to say that if these drawbacks in pitfall-trapped samples remain unresolved, this brings into question any analysis using assemblage data, such as ordination techniques, diversity indices, the determination of dominance structure and any ecological analysis or testing of theory.
While the sharpening of research questions before starting trapping is a salutary piece of advice, which will also influence the type and arrangement of traps, some problems associated with pitfall traps for general carabid beetle studies have reached a general consensus. Several of the aspects below are, however, still ignored but could be easily fixed. These include that (i) an odourless preservative is preferred, because formalin, for example, seems to attract some species and repel others (Thiele 1977); (ii) the traps should have a cover to prevent flooding, desiccation, scavenging and bycatch – a funnel to prevent escape and reduce bycatch also helps (Lange et al. 2011); (iii) traps should preferably not be used solitarily, but placed in series of at least three to five traps at distances of less than 10 m apart in order to optimise the catch and to overcome occasional trap losses; (iv) distances between sampling plots (single traps or trap groups) should be large enough to allow for sample independence (this distance will, of course, depend on the dispersal power of the focal species, see e.g. Digweed et al. 1995); and (v) the question of missing samples that inevitably occur when large numbers of traps are used over long time periods (see below). Important challenges that await study and resolution are: (i) that trap numbers and length of the trapping period do not contribute equally to the catch (Lövei and Magura 2011); (ii) how to reliably minimise the impact of trapping on assemblages and protected species (the methods of partial seasonal samples and pulsating samples, for example, have been suggested: Sapia et al. 2005); and (iii) the challenge of non-destructive carabid sampling (Bowie and Frampton 2004), such as radiotelemetry (see Negro et al. 2008).
The arrangement of pitfall traps in the field depends on the research question asked. The most popular research questions include: (i) Faunistic investigations intended to obtain an accurate species list of a given area. Here many pitfall traps should be used, also along gradients and at biotope edges; it seems that the spatial aspect is more important than the temporal one, i.e. it is better to have many traps for shorter periods of time than fewer traps for longer time periods (Lövei and Magura 2011). (ii) Community or gradient studies intended to investigate the (typical) fauna of different biotopes or at different positions along a gradient. In this case series of traps per biotope or gradient position can be used (Fig. 3a) with independent replicates (with sufficient distances between the series, see above). An example of this is the Globenet project (Niemelä et al. 2002). In some cases a row design with repeats will generate more precise information (Fig. 3b), especially when short-term movements of species along gradients are expected. The same holds for different treatments in an experimental design, such as (iii) Biological studies investigating e.g. the periodicity of one or more species within a year, to be eventually compared with different biotopes or years in phenological and/or climate studies (e.g. do species reproduce earlier or later during warmer periods or in different biotopes?). In the case of (iv) Biological studies investigating diurnal rhythms or movements of adults and larvae, a grid or matrix design (Fig. 3c) is recommended; and (v) Population studies intended to investigate the response of populations to biotic and abiotic environmental factors. Here, estimates of population densities are required and, as such, pitfall-trapped data need to be interpreted with caution.
Figure 3.
Examples of pitfall trap placements across a forest edge.
The fact that pitfall catches are a function of the species’ true population size and its activity (activity-density: Greenslade 1964; Thomas et al. 1998), creates specific statistical problems. Continuous sampling over the whole activity period can cause a potentially serious problem when the catch is analysed. Trap losses can occur at any time during the activity period and have traditionally been dealt with by standardising the catch to 100 trapping days without taking into account variability in activity across the season (Kotze and Niemelä 2002; Niemelä et al. 2002). For example, some species are more active in the spring or autumn (see Life history strategies and rhythms above), while others are active throughout the summer months. As such, a trap lost at the beginning of the continuous sampling period will have a different effect on the estimated activity-density of a spring-active species, for example, than if the trap is lost at a later stage when activity is low. When the research question involves study of the response of separate species to an environmental gradient, statistical models in which seasonality (or visit) is added as a free factor and sampling effort (number of trapping days per visit) as an offset term, and in which the response variable is specified as following a negative binomial distribution, seem to correct for seasonality and trap losses appropriately.
The reason for specifying activity-density (or abundance) data as following a negative binomial distribution (and not a Gaussian distribution, as is often done) is that ecological field data (here counts of individuals or species) seldom follow the assumptions of classical parametric statistics (Dalthorp 2004). Carabid beetles (both in terms of abundance and species) are often aggregated in space (Niemelä et al. 1986, 1992; Thomas et al. 1998) and sampling them is likely to produce an expected variance that is greater than the expected mean. Such ‘clumped’ counts data appear to be most appropriately analysed by models that incorporate extra variation, such as the negative binomial distribution (see White and Bennetts 1996; Dalthorp 2004), or quasi-Poisson methods (Ver Hoef and Boveng 2007; e.g. Elek et al. 2010). Another important advantage of using methods designed for dealing with count data (negative binomial, Poisson) is that the response variable (number of individuals or species) does not need to be transformed to comply with the assumptions of parametric test statistics, such as analysis of variance, t-test or linear regression. Surprisingly, abundance and species richness data are often log-transformed for subsequent use in parametric test procedures, even though textbooks on statistical methods in ecology (Sokal and Rohlf 1995; Crawley 2003) recommend the use of the square-root transformation to normalise count data. Nevertheless, neither square root nor log-transformed count data (for use in parametric tests) performed as well as non-transformed data (for use in a negative binomial model) (O’Hara and Kotze 2010). A possible reason for this is that count data often contain many zero values, which have to be fudged (when a log transformation is performed) by adding 0.1 or 1 to every observation – which may have unforeseen effects on estimates.
Another problem occurs when the activity density results for different species are compared. Since each species reacts differently to pitfall traps, their “catchability” will also differ, subsequently with more or less incomparable results between species. A possible solution, suggested by Den Boer, is to standardise the catches per species over the sampling sites (Turin et al. 1991). After standardisation, with the obtained “relative abundances”, multivariate methods (calculating (dis)similarities, clustering and ordination) can be used to analyse the data. Similar classifications have been carried out for Britain (Luff et al. 1989; Eyre and Luff, 1990; Mccracken 1994; Anderson et al. 2000). Although the approach of correcting and standardising the data was quite different from the Dutch method, the results for classification of the carabid habitats in the Netherlands and Britain were very similar. A study of the carabid fauna of Trento, Italy (Bonavita and Chemini 1996) in a deviating trans-alpine fauna, revealed highly corresponding results for the classification of the 48 (out of 57) species common to Italy and northern Europe. A relatively simple and flexible method developed by Dufrêne and Legendre (1997) to classify a Belgian dataset (the IndVal procedure), has the advantage that it is insensitive to the relative abundances of species. We contend that the problems associated with the comparison of assemblages sampled by pitfall trapping are still not fully resolved, but the above confirm that this method has merit in many types of investigations.
5 Population dynamics and long-term research
Since the 1960s the population dynamics of carabid beetles has been subject to the study of population persistence. During this time, discussion has revolved around how the size of populations and their fluctuations have been established, resulting in two popular theories. The first theory postulates that population sizes are balanced within narrow limits by density dependent processes, a feedback mechanism in which predators, parasites, competitors for food and other biotic aspects of the environment are involved, resulting in the regulation of population size (see Nicholson 1958). The second theory argues that the founding and re-founding of local populations take place, driven by dispersal, small population size and extinction, heterogeneity of the environment, the distinction between local (sub) and natural (entire) populations, and the genetic plasticity of species in relation to different components of the environment and to fluctuations of population size (Andrewartha and Birch 1954). Den Boer tested the latter theory by using carabid beetles as a model group. In 1959 he started pitfall trapping at several locations in the Dwingelderveld, a large area of heathland in the Netherlands, which he regarded as home to large natural populations of several carabid species. This founding/re-founding theory, the concept of metapopulation, states that natural populations consist of many local populations or colonies. Indeed, Den Boer was able to show that in a large area many local populations or interacting groups of carabids fluctuated in numbers of individuals in space and time. From these results the “spreading of risk” theory was derived (Den Boer 1968, see European Carabidologists’ Meetings (ECMs) above).
The significance of dispersal in founding, re-founding and establishment of populations was confirmed during the first ECM (see Introduction). However, the role of density dependent processes was not resolved. In 1970 in Oosterbeek, the Netherlands, an entire symposium on the Dynamics of Populations (Den Boer and Gradwell 1971) was devoted to whether or not populations were regulated. Some contributors showed examples in which density-dependent processes seemed to govern the abundance of a species, whereas others showed the opposite, so the discussion continued. Later on, again using carabid beetles, several studies were conducted to test the density dependence hypothesis. For instance Baars and Van Dijk (1984) were able to show that the number of eggs in the ovaries of females was negatively correlated with the mean density around pitfall traps. However, later on Van Dijk and Den Boer (1992) demonstrated that egg and larval mortality were too high to compensate for egg production. It was concluded that the density dependent relationship could hardly play an important role in the dynamics of the populations of Calathus melanocephalus, as shown by Baars and Van Dijk (1984). In Pterostichus oblongopunctatus the amount of food available affects the number of eggs laid. Heessen (1981) suggested that this would regulate population dynamics of this species. However, Szyszko (1981) and Den Boer (1986) observed that population explosions of certain prey species lead to a strong decline in some carabids. Vermeulen and Szyszko (1992) were able to show that in order to maintain a high level of egg production, Pterostichus oblongopunctatus has to switch prey. Presumably the right mixture of amino acids and the quality of nitrogen (White 1993) are essential for a high level of egg production. Another study on regulation in carabid populations was carried out by Brunsting et al. (1986). They showed that cannibalism occurs between larvae of Pterostichus oblongopunctatus and suggested that this phenomenon would regulate population size. However, Vermeulen (1986) could not find differences in raising Pterostichus oblongopunctatus under circumstances in which cannibalism was included and excluded, suggesting that larvae may not be actively searching for other larvae of the same species to feed upon. Cannibalism might take place only under extremely high, unnatural densities. Also, the role of competition in the population dynamics of carabid beetles has not been convincingly demonstrated so far. For example, Loreau (1990) found only weak evidence for competitive regulation in Abax ater populations. He suggested that competition might only be significant in dominant species. On the other hand, (Den Boer (1980, 1985) and Niemelä (1993) showed that competition hardly plays any role in determining population size. The discussion on whether or not regulation plays an important role in population dynamics led to a second symposium on population dynamics, this time held in Poland in 1992 (Den Boer et al. 1993). However, again only a discussion for and against regulation resulted. After this meeting the subject quickly went out of fashion and was not discussed in this way again. In 1996, Den Boer and Reddingius wrote a book in which they reviewed all the population dynamic theories so far.
At present, it is generally accepted that the persistence of carabid populations depends on the availability of sufficient suitable habitat over long periods of time, as well as on habitat quality. The latter was nicely illustrated in the Dwingelderveld, the Netherlands. These heathlands have nitrified from the 1970s onwards, associated with an almost complete disappearance of Carabus nitens there. A few years after the removal of the nutrient-rich topsoil layer by sod cutting, however, this species was again recorded in high numbers (Van Essen 1993). A similar recovery is now seen in the Mantingerveld, the Netherlands, for the same species since 2007 (Rikjan Vermeulen, pers. obs.). Because of the turnover in local populations, dispersal is necessary for a given species to (re)colonise areas where habitat patches are small. The classical technique for investigating these processes has been mark and recapture. Using modern techniques, e.g. simulation programmes (Persigehl et al. 2004) and genetic techniques (e.g. Drees et al. 2011), the relationships between populations can be demonstrated more easily.
Permanently set-up pitfall traps give an impression of the activity of different species during different seasons and between years, and produce relative estimates of population fluctuations for a particular species. In 1959, several series of permanent pitfall traps were initiated in the Dwingelderveld and later, in 1963, in the Mantingerveld (Den Boer and Van Dijk 1994). Carabid beetles from these series were collected on a weekly basis. Results from the first 6–7 years showed considerable fluctuations in the total number of individuals of a particular species collected per series between successive years. This fluctuating pattern was also different between each separate catching series within an area in the same year. These observations of asynchronous fluctuations in catches of a particular species were instrumental in the development of the “spreading of risk” theory by Piet den Boer (see above). Environmental conditions since the establishment of these series also changed. At the end of the 1960s the ground-water table gradually receded and during the 1970s the effects of air pollution became apparent: increasing acidification and eutrophication of the upper soil layers and the subsequent replacement of both Calluna and Erica by grasses. At the end of the 1980s, the local nature management authority started to artificially raise the water table, which subsequently reached its pre-1960s level during 2010–2011. At the same time the grassy vegetation, together with the polluted top soil layer, was removed by sod cutting, and grazing by cows and sheep has subsequently been introduced. Moreover, the average temperature of the area had increased by 1 oC in the last few decades. Both the increase in temperature and the hours of sunshine appear to be significant from 1988 onwards (Prins et al. 2007).
Since the establishment of these series of pitfall traps, the composition of the carabid beetle fauna has changed continuously. In the beginning of the 1970s species such as Agonum krynickii, Carabus cancellatus, Cicindela sylvatica and Cicindela germanica disappeared completely from the catches, followed by Amara quenseli and Amara praetermissa. During the same period, species such as Carabus nitens, Harpalus solitaris and Amara infima decreased significantly in numbers. The climate did not change significantly during this period, and it can be speculated that changes in the environment, as mentioned above, and habitat fragmentation (in the case of Hullenzand, Mantingerveld) may be responsible for these local extinctions and changes in population numbers. From the end of the 1990s, species such as Agonum ericeti, Cymindis vaporariorum and Cymindis macularis disappeared from the catches. This may be a consequence of climate change, since during this period environmental conditions in the heathlands improved. This is well illustrated for Carabus nitens, which became rather abundant during this period, as well as for Carabus arvensis, Nebria salina and Harpalus solitaris. From 1990 to 2004, ten species not previously recorded from these areas have been collected (Vermeulen et al. 2004). Recently, two records of Agonum viridicupreum can be added to this list (Rikjan Vermeulen pers. obs.). Apart from one, all of these newly recorded species have their center of distribution south of the Netherlands, suggesting that their appearance is related to climate warming. Similarly, the virtual disappearance of the northern species, Agonum ericeti, may be related to this phenomenon. Adequate management may, to a limited extent, compensate for the effects of climate change. The northerly distributed Carabus nitens that almost disappeared from both the Mantinger- and Dwingelderveld, made a rapid comeback after top-soil removal and sod-cutting.
However, the dramatic decline and extinction of the highly hygrophylic Carabus clatratus in Italy may not be entirely related to climate change. Carabus clatratus is one of the most localised and endangered carabid species in Europe, and its disappearance from Italy, and possibly also France, is possibly a consequence of the colonisation of its wet biotopes by the alien red swamp crayfish, Procambarus clarkii, which preys on adults of Carabus clatratus (Casale and Busato 2008).
Long-term data on weekly catches can also be used to monitor phenological changes in species. For example, compared to the period prior to 1988, the activity of Amara equestris, Carabus arvensis, Poecilus lepidus and Poecilus versicolor started earlier in the season.
As far as the consequences of climate change, management and other environmental changes are concerned, it is of great importance to continue long-term observational studies of carabid beetles, such as that in Drenthe, so that future changes can be monitored and possibly explained. Such long-term sampling programmes for carabid beetles are also known from Poland, Germany and Italy.
6 Bioindicators
Carabids are excellent model organisms for research on ecological and conservation theory. These beetles readily respond to abiotic and biotic variation, and to disturbances and management (e.g. Lövei and Sunderland 1996; Rainio and Niemelä 2003). This evidence has led many to suggest carabids to function as ‘indicators’. An indicator is a taxon or a structure “whose characteristics (...) are used as an index of attributes too difficult, inconvenient, or expensive to measure for other species or environmental conditions of interest” (Landres et al. 1988). However, using this definition many, if not most, of carabid ‘indicator’ studies appear to only demonstrate individualistic responses to environmental variation. But instead of investing resources for finding new indicator taxa, environmental managers should test and select taxa that are already well known and easily sampled, and that cover multiple dimensions of biodiversity (Taylor and Doran 2001), and critically evaluate their indicator functioning (Langor and Spence 2006). Carabids fulfil the former but the latter aspect requires further attention.
European carabids have certain qualities that make them good candidates for indicators. They are taxonomically well known, with relatively stable systematics, and their ecology has been widely studied (Lövei and Sunderland 1996). Variation in carabid morphology, life history strategies and small-scale abiotic and biotic requirements are extensively documented (e.g. Lindroth 1961–1969, 1985, 1986). Carabids also respond predictably to not only small-scale but also to landscape- and even continent-level phenomena (e.g. Hengeveld 1987; Kotze and O’Hara 2003; Koivula and Spence 2006). Moreover, they are relatively easy to collect in high numbers using standard methods. But can carabids reflect environmental variation in ways useful for conservation assessment purposes? Knowledge of carabid indicator functioning, using the categories listed in Lindenmayer et al. (2000), is briefly summarised below (see Koivula 2011 for a complete evaluation).
i. Taxon indicators. The presence of a taxon indicator reflects the presence of a set of other species, and its absence indicates the lack of the entire set of species. Perfect multi-taxon richness overlaps may be rare (e.g. Jonsson and Jonsell 1999; Sætersdal et al. 2005; Similä et al. 2006), which highlights the importance of using multiple taxa in environmental assessments (Taylor and Doran 2001; Duelli and Obrist 2003). Carabid functioning as taxon indicators mostly relies on weak correlations among taxa.
ii. Keystone indicators. These species affect their environment disproportionately strongly relative to their abundance. In field and laboratory conditions, carabids forage on slugs and pest insects (e.g. Kromp 1999). Hance (1987) showed that, using enclosures with different carabid densities, carabids have the potential to significantly prey on pest insects foraging on crop plants with economic benefits.
iii. Pollution indicators. These taxa reflect human-altered abiotic conditions. Heavy metals in the soil negatively affect carabids (e.g. Maryański et al. 2002; Ermakov 2004), and in agro-ecosystems, pesticides and fertilizers affect carabids, at least in the short term (e.g. Huusela-Veistola 1996; Kromp 1999).
iv. Dominant indicators. These taxa make up much of the total biomass or the number of individuals in an area of interest and predict particular ecosystems or assemblages. Many common carabid species are succession and habitat-type generalists (Lindroth 1985, 1986; Niemelä et al. 2007), so their numbers may not indicate aspects useful for conservation or management. Mean Individual Biomass (MIB), on the other hand, links carabid biomass to succession without considering species entities (Szyszko et al. 2000). However, the ‘behaviour’ of MIB along succession should be examined in detail before applying it in conservation and management.
v. Environmental indicators. These should reliably reflect particular environmental conditions. Although carabids have the potential to reflect soils, wetness and habitat-type variation (e.g. Thiele 1977; Lindroth 1985, 1986), they cannot currently compete with plants as indicators of these factors.
vi. Early-warning signallers (true bio-indicators). These taxa are extremely sensitive to changing environmental conditions. Carabid evidence is scarce, but some carabids have apparently undergone shifts of tens of metres in altitude over 10–20 years (Assmann 2009; Pizzolotto 2009, David Kavanaugh, pers. comm.), coinciding with climate warming (Parry et al. 2007, see Population dynamics and long-term research above). These observations suggest good potential in, for example, climate-change and urban-spread research.
vii. Disturbance indicators. These taxa reflect natural and human-caused disturbances. Carabids readily respond to agriculture and forestry (for reviews, see Lövei and Sunderland 1996; Kromp 1999; Niemelä et al. 2007). Their indicator functioning may hold at a general level: they respond similarly to environmental change as many other taxa do (e.g. Barbaro et al. 2005). But indicators should not be used for self-evident patterns: the ecological impact of clear-cutting, for example, does not require an indicator.
Clearly, carabids have good potential for becoming useful indicators for conservationists and environmental managers. Certain obstacles still need to be overcome. First, the functioning and accuracy of carabids to predict habitats or species requiring conservation action should be critically evaluated. According to the indicator definition of Landres et al. (1988), none of the above examples indicate that carabids function as particularly useful indicators. Thus, for a conservationist, carabid responses should be considered as individualistic as long as there is no evidence for their responses to reliably predict responses of threatened taxa or particular, difficult-to-observe conditions. This is important because there is very little room for error if threatened species or habitats are at stake. Strict tests must thus be applied to evaluate indicator functioning (Langor and Spence 2006). Second, the relationship between carabid responses and other taxa should be considerably clarified (Rainio and Niemelä 2003) before using these beetles in environmental assessments. Third, it is unclear whether carabids reflect aspects not attainable using other indicators (apart from their individualistic response) and whether conditions exist under which carabids really are the most cost-efficient indicator taxon. Currently widely used, easy-to-use, relatively cheap and economic tools for assessing the state of the environment include vegetation, habitat structural elements, satellite and aerial photos, as well as weather and land-use inventory data.
The focus of carabidologists should perhaps be changed from total species richness to the indicator potential of single species, groups of specialists or functional groups. We lack an explicitly defined ‘niche’ of these beetles in environmental assessment protocols. Cases for carabids fulfilling the conservationists’ definition for a useful indicator (Landres et al. 1988) will possibly be documented in the near future, but their indicator functioning may always remain context specific.
7 Carabid conservation, protection and habitat management
Conservation may mean protecting particular species or patches of habitat against alteration, generally human-caused, but the term may also include operations characterised by an active human role (e.g. Freitag and Kavanaugh 1993; Den Boer and Van Dijk 1994; Sutherland 1998; Gaston and Spicer 2004). Examples include the maintenance of areas of high natural value, the restoration of patches to a state they are presumed to once have represented (often referred to as ‘natural’ state), and the artificial conversion of one habitat type to another. The latter may be required in landscapes where habitat for a threatened species has become rare (see Negro et al. 2008) and new habitat patches are unlikely to appear through natural processes. Such cases might be found, for example, within urban areas. These active operations of patch maintenance, restoration and creation are collectively called ‘conservation management’.
Insect conservation management is a relatively new research discipline, both generally and in the context of carabid beetles (e.g. Lewis et al. 2007; Leather et al. 2008; New 2010). The restoration and artificial creation of habitats − two elements of conservation management − have been important components of carabid conservation since the 1980s (e.g. Thomas 1990; Främbs 1990; Blake et al. 1996). Conservation became an important topic for the ECMs since the Hungarian meeting in 1986. Before that meeting, conservation issues were only occasionally discussed, but from then on, both conservation in general (e.g. identifying diversity hotspots and gathering data on endemic and rare species) and practical conservation management in particular have been among key topics and have altogether consistently made up over 20% of papers in the proceedings. Generally, almost any piece of knowledge on carabid ecology can be applied in conservation-management policy and action to support these beetles and associated epigaeic fauna. In Europe and North America, information necessary for efficient conservation − on carabid ecology and threats − is readily available (Maelfait et al. 1994; Lövei and Sunderland 1996; see also national lists of threatened species). However, the functioning of active management for the benefit of threatened carabid species urgently demands critical evaluation and detailed information. For instance, according to Desender et al. (2010), the decline of carabid beetles in Belgium between the period <1950 and 1950–1985, had halted for a considerable number of species. During the period 1986–2008, however, 60% of these species still had not reached the same distribution area as in the first half of the 20th century, notwithstanding many initiatives and large scale active management. Most of these species now only occur in large and high-quality nature reserves with the last remnants of semi-natural biotopes and have, at present, little or no possibilities to further increase their distribution range.
Here the advances in conservation management, mostly as derived from the proceedings of the previous ECMs are discussed under four topics: (i) Which species characteristics are particularly associated with threatened species? (ii) In which habitat types can conservation of carabids best be realised? (iii) What do we know about habitat connectivity as a way to conserve carabids? (iv) How does conservation management of habitats affect carabids?
i. Ecological and habitat characteristics of threatened species. To study which ecological and habitat characteristics of carabids are associated with species being threatened, national species lists and their IUCN categories for five countries are used as examples: Belgium, Sweden, Denmark, Norway and Finland (respectively Desender et al. 2008a, b; Gärdenfors 2005; Pedersen and Wind 2009; Kålås et al. 2006; Rassi et al. 2001). This dataset is complemented with four regional lists of threatened species from Niedersachsen and Bremen, Germany; Nordrhein-Westfalen, Germany; Wadden Sea area; and a preliminary red list for Drenthe, the Netherlands (respectively Assmann et al. 2002; Schüle and Terlutter 1998; Mahler et al. 1996; Noordijk and Vermeulen 2009). For analytical purposes, species characteristics were collected from Lindroth (1985, 1986), Desender (1986), Turin and Den Boer (1988), Desender and Turin (1989), Turin (2000), Anonymous (2006) and Desender et al. (2008a). Several characteristics were evaluated, such as the roles of body size, wing morphology, and associations with shadiness and moisture. This evaluation was done by calculating percentages per size, wing morphology, shadiness and moisture classes for all species (for the five countries), for species classified as threatened by IUCN categories NT (Near Threatened), VU (VUlnerable), EN (ENdangered), CR (CRitically endangered) and EW (Extinct in the Wild; also RE, i.e. Regionally Extinct, in some national lists), and also the proportion of threatened species over all species within a given class. Occasionally, certain information for some species was lacking and these were (partly) removed from the analysis. For example, if for a certain species information was unavailable on wing morphology, it was omitted from the wing morphology analysis but retained in other analyses.
Carabids mostly fell into mid-size classes (43–46% of all species were 4.1–8.0 mm and 28–31% were 8.1–16.0 mm), were macropterous (64–71%) and were associated with open areas (63–64%), but were quite evenly distributed among moisture-association classes (see columns “All” in Table 3). Carabids classified as being threatened roughly complied with these figures (columns “IUCN” in Table 3): also these species were mostly mid-sized (26–50% were 4.1–8.0 mm and 24–39% were 8.1–16.0 mm), macropterous (64–81%) and open-area associated (63–79%; very shady habitats had only 2–12%). However, threatened species were more often associated with either very wet (34–53%) or very dry habitats (32–47%) than with “average” or moist/dryish conditions (12–30%). This dichotomous association with both very dry and very wet habitats was much more pronounced in the four Nordic countries and in the two areas in Germany than in Belgium or in Drenthe (Table 3).
Table 3.
Morphological and habitat-association characteristics of all carabid species found in a given area (“All” columns; % of species), of species classified as threatened according to the IUCN (“IUCN”; categories NT, EN, VU, CR and EW pooled; % of species), and proportion of threatened species of all species within a given category (“% IUCN”). For example, the value “50” for BEL % IUCN >16 mm indicates that in Belgium, of all species with body size >16 mm, 50% are considered threatened. Values for “All“ and “IUCN“ columns make up 100% for each area/country. BEL = Belgium; SWE = Sweden; DEN = Denmark; NOR = Norway; FIN = Finland; Niede = Niedersachsen and Bremen, Germany; Nordr = Nordrhein-Westfalen, Germany; Wadde = Wadden Sea area; and Drent = Drenthe, the Netherlands (proposed Red Data list). For the last four areas, only species classified as threatened according to the IUCN are shown. For species and their characteristics data, see text.
| Classes | BEL | SWE | DEN | |||||||
|---|---|---|---|---|---|---|---|---|---|---|
| All | IUCN | % IUCN | All | IUCN | % IUCN | All | IUCN | % IUCN | ||
| Body size | ||||||||||
| 0.1-4.0 mm | 18 | 13 | 18 | 19 | 16 | 14 | 19 | 9 | 13 | |
| 4.1-8.0 mm | 44 | 40 | 25 | 44 | 31 | 11 | 43 | 38 | 25 | |
| 8.1-16.0 mm | 31 | 35 | 31 | 29 | 40 | 23 | 30 | 41 | 39 | |
| >16 mm | 7 | 12 | 50 | 8 | 13 | 28 | 8 | 12 | 46 | |
| Wing morphology | ||||||||||
| Macropt | 71 | 69 | 23 | 68 | 69 | 15 | 66 | 68 | 29 | |
| Poly/dimo | 16 | 18 | 26 | 19 | 19 | 14 | 14 | 16 | 33 | |
| Brachypt | 13 | 13 | 23 | 13 | 12 | 13 | 20 | 16 | 23 | |
| Shadiness | ||||||||||
| Shady (forest) | 11 | 8 | 24 | 11 | 11 | 17 | 11 | 10 | 25 | |
| Generalist | 26 | 23 | 29 | 25 | 11 | 7 | 25 | 26 | 29 | |
| Open | 63 | 69 | 36 | 64 | 78 | 20 | 64 | 64 | 28 | |
| Moisture | ||||||||||
| Water/wet | 39 | 35 | 29 | 38 | 35 | 15 | 37 | 42 | 32 | |
| Moist-dryish | 29 | 23 | 26 | 30 | 18 | 10 | 31 | 22 | 20 | |
| Dry | 32 | 42 | 43 | 32 | 47 | 24 | 32 | 36 | 30 | |
| NOR | FIN | Niede | Nordr | Wadde | Drent | |||||
| Classes | All | IUCN | % IUCN | All | IUCN | % IUCN | IUCN | IUCN | IUCN | IUCN |
| Body size | ||||||||||
| 0.1-4.0 mm | 18 | 13 | 13 | 18 | 15 | 10 | 19 | 17 | 19 | 16 |
| 4.1-8.0 mm | 46 | 38 | 15 | 45 | 26 | 7 | 44 | 40 | 50 | 33 |
| 8.1-16.0 mm | 28 | 36 | 24 | 31 | 53 | 21 | 30 | 34 | 24 | 39 |
| >16 mm | 8 | 13 | 30 | 6 | 6 | 12 | 7 | 9 | 7 | 12 |
| Wing morphology | ||||||||||
| Macropt | 64 | 63 | 16 | 70 | 81 | 12 | 71 | 74 | 74 | 64 |
| Poly/dimo | 21 | 16 | 13 | 17 | 15 | 9 | 17 | 14 | 21 | 21 |
| Brachypt | 15 | 21 | 23 | 13 | 4 | 3 | 12 | 12 | 5 | 15 |
| Shadiness | ||||||||||
| Shady (forest) | 10 | 9 | 15 | 10 | 3 | 4 | 6 | 7 | 2 | 12 |
| Generalist | 26 | 21 | 15 | 27 | 21 | 9 | 28 | 30 | 19 | 21 |
| Open | 64 | 70 | 20 | 63 | 76 | 15 | 66 | 63 | 79 | 67 |
| Moisture | ||||||||||
| Water/wet | 37 | 47 | 22 | 40 | 47 | 14 | 46 | 51 | 53 | 31 |
| Moist-dryish | 32 | 17 | 10 | 30 | 12 | 5 | 16 | 17 | 14 | 30 |
| Dry | 31 | 36 | 20 | 30 | 41 | 17 | 38 | 32 | 33 | 39 |
The proportion of threatened species over all species in the five countries revealed some important issues (columns “% IUCN” in Table 3). First of all, relative to the total number of species per category, larger species tended to be more often threatened than smaller species (see Kotze and O’Hara 2003). For size classes 8.1–16.0 mm and >16 mm, the proportions of threatened species were 21–39% and 12–50%, respectively, whereas for the size classes 0.1–4.0 mm and 4.1–8.0 mm, they were 10–18% and 7–25%, respectively. After pooling species into larger (>8.1 mm) and smaller (0.1–8.0 mm) size classes, proportions of these were between 19–40% (mean 28%) and 8–23% (mean 16%), respectively. Regarding wing morphology, the proportions of threatened species were rather even among the categories. In this respect, wing morphology was not clearly related to species being threatened, except for the slight tendency of wing-polymorphic species being proportionally more frequently threatened in Denmark and brachypterous species in Norway. Regarding shadiness associations, open-area species included proportionally slightly more threatened species than did species of very shady habitats or shadiness generalists. Regarding moisture associations, species associated with very dry habitats included proportionally more threatened species (17–43%) than wet-habitat species (14–29%) or moist/dryish-habitat species (5–26%).
To the extent one can generalise from these figures, in northern and western Europe (see also Casale and Busato 2008 for southern Europe) particular attention should be paid to large carabids, species associated with very dry, open habitats (e.g. sand dunes, heathlands and calcareous meadows; see national Red Lists) and water-associated species (e.g. freshwater stream specialists and salt-marsh species; see national Red Lists). However, as is evident from the variation in percentages presented in Table 3, particular targets of conservation and management (habitat types and species) should vary from one area to another. Below, research-based evidence on how to protect these carabids by the application of conservation management is reviewed.
ii. Habitat selection for conservation efforts. Undisturbed mature ecosystems, particularly nature reserves, are vital for the conservation of many carabid species (e.g. Desender 2005; Skłodowski 2006). Also edge habitats and habitat mosaics may be important for carabid conservation (e.g. Kotze 2000; Falke et al. 2000; Hatteland et al. 2005; Andorkó and Kádár 2006). The scarcity of certain habitat types has increased the need for active maintenance of remaining patches. For example, Bérces et al. (2008) used field and museum data, original research and communication among entomologists, and showed that Carabus hungaricus can best be protected by active management of open meadows. Due to the on-going loss of natural and semi-natural areas and the intensification of agricultural practices in many countries, also particular anthropogenic habitats have become important for carabid conservation. Examples include roadside verges, former agricultural fields, urban waste-grounds, and sand and gravel pits (Plachter 1986; Eversham et al. 1996; Telfer and Eversham 1996; Schwerk 2000; Versteirt et al. 2002; Koivula and Kotze 2005).
iii. Habitat connectivity. Habitat-patch isolation and fragmentation may be of major concern for carabids (De Vries 1994; Kinnunen et al. 1996; Noordijk et al. 2006; Hendrickx et al. 2009). A number of means of reducing the impact of fragmentation have been suggested (Vermeulen et al. 2002). For forested environments, Terlutter (1990) discovered for Carabus auronitens two different gene flows from two old forest remnants into a recent, regenerated forest-field mosaic. Later on, Petit (1994) underlined the importance of hedgerow networks for forest carabid assemblages. However, more recent hedges may be sub-optimal for this purpose (Thiele 1971; Gruttke 1994). For open areas, on the other hand, Vermeulen and Opsteeg (1994) showed that roadside verges might be used either as habitat or as movement corridors connecting heathland patches. Both purposes may be served by roadsides, as shown by (Koivula (2002a, 2005) for Finnish forest roads and Noordijk (2009) for highway verges in the Netherlands. Moreover, Vermeulen and Spee (2005) stressed the importance of source habitats for nature restoration sites. Hence, the remaining patches of natural habitat, corridors of similar, often man-made environments, and artificially created, larger patches may together form an efficient patch network for carabid conservation.
iv. Habitat management. Because many important natural processes (wildfire, flooding, wind, grazing and insect outbreaks) are effectively prevented in many areas, particularly in urban environments, active maintenance is considered necessary to preserve certain vegetation types. Carabids respond varyingly to these efforts (Versteirt et al. 2002; Cuesta et al. 2006; Taboada et al. 2006a). The effects of grassland management were discussed in depth by Rushton et al. (1990) who found that some species avoid intensively managed sites, some are favoured by these, while others showed intermediate or no detectable responses. Similarly, Blake et al. (1996) showed that vegetation management in wildflower meadows resulted in a decrease in large species and an increase in xerophilous species, while species characteristic of areas with ‘natural’ conditions were absent. Like mowing, grazing also profoundly affects carabids: its intensity determines assemblage composition (McFerran et al. 1994). Cole et al. (2006) showed that intensive grazing decreases the abundance of large Carabus species more than less intensive grazing. These studies indicate that variation in management leads to variation in carabid beetle assemblages. In riparian environments, Fuellhaas (2000) showed that raising the water-table level increases the number of hygrophilic species. Främbs (1990) studied regenerating peat bogs and found that although carabid diversity increased, the peat-bog specialist Agonum ericeti remained absent. Drees et al. (2007) argued that this might be related to habitat quality, in this case the lack of peat-producing vegetation.
To summarise, (i) Carabid conservation should give special attention to very large species, and species associated with both very wet and very dry, exposed conditions, (ii) Old and undisturbed natural areas are important for many specialists, but conservation of pioneer or open-habitat species can be realised in many anthropogenic areas as well (Fig. 4), (iii) Fragmentation potentially isolates local populations, but its effects can be decreased by maintaining large, inter-connected areas, corridor networks, and designing restoration areas near potential source areas, and (iv) Guidelines for active management of carabid habitats are difficult to draft, as some species respond negatively to any disturbance, including conservation management. However, many species urgently need small-scale management that keeps habitats constantly at some preferred successional phase; most of these species are subject to severe stress in modern, fragmented landscapes.
Figure 4.
Cylindera germanica (Photo by Jinze Noordijk)
The conservation of carabids and their habitats is far from perfect. This issue is complicated by the fact that, due to these beetles’ mobility, occupation of varyingly sized habitat patches, varying degrees of specialisation, and development through numerous developmental phases, their ecological requirements vary in time and place. As habitat patches of carabid assemblages usually include several vegetation types and/or physical structures, a conservation approach targeted for maintaining only particular vegetation types or high plant diversity may not always be appropriate for the conservation of arthropod assemblages (Panzer and Schwartz 1998; Dennis et al. 2007). Moreover, the high number of carabid species, each with specific demands, makes it difficult to define a single conservation strategy. Protection of whole landscapes with mosaics of distinct habitat types may prove efficient for carabid conservation. Simultaneously, the natural variety of successional stages should be conserved, as particular stages can be crucial for certain species (cf. Niemelä et al. 2007). Moreover, some species – such as Amara plebeja (van Huizen 1977) – possibly require more than just one habitat type and/or successional stage to persist in a landscape (‘landscape species’; Szyszko 2004; Szyszko et al. 2011; Axel Schwerk, pers. comm.). Habitat patches within these mosaics should include particular structures, such as micro-relief, patches of bare sand, stony patches, small water bodies, heaps of decaying plant material, and dead wood, features that are often of no special importance for plants and vertebrates and therefore often ignored if conservation management is based on vegetation data alone. Thus, a broad landscape approach, supplemented by these small-scale structures, may produce good results for the conservation of carabid beetles (Kirby 1992; New 1995, 2010; Samways 2005, 2007; Haslett 2007).
8 Landscape ecology
How carabid beetles perceive space may influence habitat selection, home ranges, the dispersal of individuals and the dynamics and distributions of populations. Furthermore, the amount, extent and spatial arrangement of suitable habitats within a landscape (i.e. landscape composition and configuration) may affect long-term population persistence. Thus, although the spatial distribution of carabid beetles may be primarily determined by microhabitat conditions and biotic interactions at the local scale, identifying general patterns of carabid responses to landscape features may help us to understand how species, functional groups and assemblages effectively distribute, and to predict how they will cope with current and future land-use and climatic changes. As such, the spatial context related to a species’ distribution patterns is an essential component when studying how global changes affect carabid species conservation.
Over the last 40 years, investigations of ground beetle landscape ecology have demonstrated that landscape features influence not only the spatial distribution of these beetles, but also their population dynamics (Matalin 1997c; Bommarco 1998) and genetic structure (Brouat et al. 2003; Keller et al. 2004; Desender et al. 2005; Sander et al. 2006). From the late 1950s until the mid 1970s, contributions to carabid beetle ecology aimed at characterising the structure and composition of communities occurring in specific types of landscapes, which were, at that stage, considered as homogeneous entities (e.g. forested vs. open landscapes; see Thiele 1977). Research developed in the 1980s and 1990s confirmed the significance of heterogeneity within landscapes and thus addressed the role of singular landscape elements or habitat types for the carabid fauna in a variety of either natural or highly-modified and simplified landscapes. In tests of the application of the theory of island biogeography (MacArthur and Wilson 1967) to carabid communities, it has been shown that local communities are not simply a passive random sample of the regional species pool, but that species are filtered according to the association of their life history traits to habitat quality, configuration and biotic interactions (e.g. Ranta and Ås 1982; Niemelä et al. 1985; De Vries et al. 1996). Studies that have looked for an island effect in carabid assemblages of patches of terrestrial habitats have generally concluded that such patches are not sufficiently isolated to represent islands, due to the strong dispersal capacity of many carabid species (Davies and Margules 1998; Magura et al. 2001; Brose 2003). Studies have also been conducted on carabid assemblages of real islands, and these too have concluded that a simple species-area relationship explains the differences in carabid species richness between islands of different size better than distance from mainland populations (Kotze and Niemelä 2002; Zalewski 2004).
In recent literature, studies on the importance of the landscape context in determining the occurrence of carabid species based on different aspects of landscape composition, configuration, connectivity, history, land-use type and intensity have proliferated (e.g. Purtauf et al. 2004; Bräuniger et al. 2010; Gardiner et al. 2010; Nabe-Nielsen et al. 2010; Woodcock et al. 2010). Many studies analysed the influence of the landscape context on overall carabid beetle activity density and species richness, often finding no statistically significant effect. Mostly, changes in landscape features have been related to shifts in carabid species composition, and variations in the activity density of individual species and ecologically meaningful groups (e.g. Niemelä 2001; Kotze and O’Hara 2003; Niemelä et al. 2007; Niemelä and Kotze 2009).
Agricultural landscapes in particular, driven by daily, seasonal and annual fluctuations, soon became the subject of many carabid beetle surveys, followed by an extensive number of publications to date (e.g. Kinnunen et al. 2001; Holland 2002). In general, the basic composition of the carabid fauna of agricultural mosaic landscapes appears to be surprisingly similar across countries (Luff 2002), dominated by eurytopic species, which are highly tolerant to disturbance. However, the size, amount, isolation and spatial arrangement of agricultural patches, the composition of the arable mosaic, as well as the occurrence of permanent landscape elements (e.g. hedgerows, field margins, natural woodlands and grasslands), affect carabid beetle assemblages (Kinnunen et al. 1996, 2001; Burel et al. 1998; Petit and Usher 1998; Fournier and Loreau 2001; Millán de la Peña et al. 2003; Aviron et al. 2005; Purtauf et al. 2005; Griffiths et al. 2007; Hendrickx et al. 2007; Saska et al. 2007).
In forest ecosystems, natural and anthropogenic disturbances create a dynamic mosaic of successional habitat patches for carabids (e.g. Bouget and Duelli 2004 for windstorm disturbance). Each forest successional stage is characterised by a specific carabid assemblage, in terms of species composition as well as ecological group composition, with the greatest differences between early and advanced stages (e.g. Szyszko 1990; Niemelä et al. 1996; Butterfield 1997; Koivula et al. 2002; Du Bus de Warnaffe and Lebrun 2004; Richard et al. 2004; Magura et al. 2006; Taboada et al. 2008). Changes in population dynamics and morphological traits also take place through succession (e.g. Szysko et al. 1996 for Pterostichus oblongopunctatus, Table 4). Changes in the carabid fauna are possibly correlated with the amount of carbon accumulation in the forest system, i.e. in the wood, litter and mineral soil (Szyszko 2010; Szyszko et al. 2011). The increase of carbon in the mineral soil is related to the decomposition of litter by the macrofauna. For pine stands in Poland, Szyszko (1986a) demonstrated that biomass of the macrofauna is correlated with parameters of the carabid fauna, such as species number and Mean Individual Biomass (MIB). MIB increases as succession progresses (Szyszko 1986b; Szyszko et al. 2000; Szyszko 2004), suggesting that this measure functions as a good indicator of the state of succession (see Bioindicators above). The rate at which species composition changes during succession and the successional trajectory followed by the carabid assemblages depends on environmental conditions, such as soil properties (Szyszko 1986b, 1990; Schwerk 2008), dominant tree species (Du Bus de Warnaffe and Lebrun 2004), and the type of disturbance that initiated the succession (Du Bus de Warnaffe and Lebrun 2004). Indeed, the larger the newly-created gap is and the fewer trees retained, the more severe the perturbation for carabid assemblages (Koivula 2002b following timber harvest; Bouget 2005 and Skłodowski and Garbalińska 2010 following windthrow gap). As a consequence, the maintenance of a variety of successional phases of the forest cycle results in increased heterogeneity at the landscape level and, therefore greater regional carabid diversity (e.g. Mullen et al. 2008; Taboada et al. 2008). Thus, the effects of forest landscape features on the carabid fauna have also been extensively addressed as regards to landscape heterogeneity, the occurrence, composition and spatial configuration of either natural or human-modified habitats (e.g. proportion of deciduous vs. coniferous forests, age and extent of exotic plantations, forest edge density and permeability), the role of particular landscape elements (e.g. retention tree groups), and the landscape context resulting from historical and/or recent management practices (Koivula et al. 2002; Bouget 2004; Barbaro et al. 2005; Matveinen-Huju et al. 2006; Taboada et al. 2006b; Barbaro et al. 2007; Niemelä et al. 2007; Fuller et al. 2008; Pawson et al. 2008; Barbaro and van Halder 2009).
Table 4.
Changes in carabid fauna, interaction groups and populations of Pterostichus oblongopunctatus with changes in habitat (according to Szyszko et al. 1996, reprinted and modified with permission from Aarhus University Press).
| Comparatively early stage of succession | Comparatively late stage of succession | |||
| Carabidae fauna | ||||
| low state of development of fauna | → | high state of development of fauna | ||
| high number of species | → | low number of species | ||
| small individuals | → | large individuals | ||
| low mean individual biomass (MIB) | → | high mean individual biomass (MIB) | ||
| Interaction group of Pterostichus oblongopunctatus | ||||
| long period of activity | → | short period of activity | ||
| long survival of adults | → | short survival of adults | ||
| complicated age structure | → | simple age structure | ||
| small individuals (imago) | → | big individuals (imago) | ||
| high proportion of males | → | high proportion of females | ||
| low number of eggs in ovaries | → | high number of eggs in ovaries | ||
| high number of eggs laid? | → | low number of eggs laid? | ||
| good food situation for adults? | → | bad food situation for adults? | ||
| bad food situation for larvae? | → | good food situation for larvae? | ||
| unable to fly? | → | able to fly? | ||
| uneconomic life strategy | → | economic life strategy | ||
| Populations of Pterostichus oblongopunctatus | ||||
| asynchronously fluctuating interaction groups | → | synchronously fluctuating interaction groups | → | asynchronously fluctuating interaction groups |
| low probability of high fluctuations of numbers | → | high probability of high fluctuations of numbers | → | low probability of high fluctuations of numbers |
| resistant population | → | not very resistant population | → | resistant population |
Much effort has been devoted to investigating how carabid beetles are distributed in fragmented landscapes and insular environments (for reviews, see Niemelä 2001 and Kotze 2008, respectively). Carabid responses to fragmentation depend on the geographical context, are species specific and, to a great extent, relate to species’ life history traits and habitat associations (e.g. Koivula and Vermeulen 2005; Gaublomme et al. 2008). In a fragmented landscape context, mobility is crucial for persistence, especially for specialist and scarce species (De Vries 1994; De Vries et al. 1996). In general, good dispersers and abundant species are expected to maintain populations in small and isolated patches through recolonisation of empty patches, whereas poor colonisers and scarce species may not be able to do so (Den Boer 1977; Niemelä 2001). Hostile types of matrix or linear elements in the landscape can act as dispersal barriers for specialist species. For instance, some forest species are reluctant to cross highways (Mader 1984; Koivula and Vermeulen 2005) or open habitats (Plat et al. 1995; Riecken and Raths 1996), while other stenotopic species effectively move along hedgerows (Burel 1989; Plat et al. 1995; Charrier et al. 1997) and roadside verges (Vermeulen 1993, 1994; Vermeulen and Opdam 1995), which function as movement corridors for such species. Dirt roads in forested landscapes may serve as dispersal corridors for open habitat species (Koivula 2002a), and roadsides overgrown with poplars have been suggested to serve as corridors for forest species with low dispersal power (Dymitryszyn et al. 2003). Attempts to improve the connectivity of landscape elements by means of corridors may have contrasting effects on different carabid species according to their habitat requirements and, hence, new approaches regarding this matter are now under evaluation, such as semi-open corridors (Eggers et al. 2010) and innovative, small scale forest harvesting techniques (Koivula et al. 2002, see also Carabid conservation, protection and habitat management above).
Further studies have investigated the responses of ground beetles to anthropogenic or human-modified landscapes and urban environments (e.g. Czechowski 1982; Klausnitzer and Richter 1983; Šustek 1987, 1992; Niemelä et al. 2002). These investigations have identified distinct sets of species associated with the urban cores or city centres (but see Niemelä et al. 2002). However, for a considerable number of these species, urban populations may be dependent on recruitment from populations in the urban periphery (Klausnitzer and Richter 1983). The possible effects of urbanisation on carabid population genetics (i.e. genetic diversity and differentiation) remain unclear (Desender et al. 2005). In general, the overall abundance and species richness of carabids decrease with increasing urbanisation (Niemelä and Kotze 2009; but see Magura et al. 2010). Also, large species tend to be relatively scarce in urban habitats, resulting in a decline in average body size in urban areas compared to less disturbed ones, both for forest assemblages (Niemelä et al. 2002; Ishitani et al. 2003; Sadler et al. 2006; Elek and Lövei 2007) and those of open habitats (Czechowski 1982; Šustek 1987; Venn 2007). Flightless species also tend to be relatively scarce in urban assemblages (Venn et al. 2003; Sadler et al. 2006). Other responses detected in carabid assemblages (either in the proportion of species or the number of individuals) to urbanisation include (i) a decrease in species with restricted geographical ranges, along with the enhancement of those distributed over broad ranges; (ii) a decline in oligotopic, stenotopic and specialist species, whilst eurytopic, polytopic and generalist ones increase; (iii) a decrease in forest species and associated increase in open habitat species; (iv) an increase in xerophilic and mesohygrophilous species at the expense of more hygrophilous species; and (v) an increase in omnivorous species and a corresponding decrease in zoophagous species. Whilst stenotopic and specialist species generally decline with increasing urbanisation, some extremely harsh urban habitats accommodate these species, such as populations of Amara equestris in central reservations of a busy ring road in Helsinki, Finland (Koivula et al. 2005). In fact, Eversham et al. (1996) reported that more than 35% of Britain’s rare and scarce carabids are to be found from manmade sites, Omophron limbatum and Dyschirius obscurus exclusively so. Subsequent to numerous urbanisation studies from single cities, the GLOBENET project (Niemelä et al. 2000; http://www.helsinki.fi/science/globenet) was established to apply a standard urbanisation gradient approach in cities across the globe (nine cities located in Europe, Japan and Canada). The main findings indicated that the carabid fauna of urban forested habitats display uniform patterns of response to the degree of urbanisation of the ‘concrete’ matrix (Niemelä and Kotze 2009; Magura et al. 2010).
In the BIOASSESS project (http://www.nbu.ac.uk/bioassess), global patterns in carabid responses to a land-use intensity gradient from old-growth or unmanaged forests to arable crop-dominated landscape across ten countries, have so far reported effects on overall species richness, number of individuals, ecological groups and species composition (Grandchamp et al. 2005; Schweiger et al. 2005; Vanbergen et al. 2005, 2010; Hendrickx et al. 2007; Martins da Silva et al. 2008). Similarly, changes in landscape structure over time (i.e. landscape history) have been addressed when investigating carabid population declines or range-size modifications in human-altered landscapes (Turin and Den Boer 1988; Desender et al. 1994b; Petit and Burel 1998; Kotze and O’Hara 2003). Additionally, these investigations have related contemporary distribution patterns of carabid endemism, rarity and habitat specialisation across landscapes to landscape history.
Even now, the spatial scale at which carabid beetles relate to resources across landscapes is not completely understood. Future studies should accomplish multiscale approaches that consider a wide range of fine and coarse grains at which each carabid species may perceive the landscape, depending on its mobility and body size (e.g. Burel et al. 2004; Aviron et al. 2005; Janssen et al. 2009). The spatial distribution of carabid species in a given landscape is nearly always aggregated at some scale (see e.g. Niemelä et al. 1996; Thomas et al. 2002), which suggests that spatial autocorrelation should always be taken into account (Barton et al. 2009). Additionally, the use of multiple habitats by carabid species in mosaic heterogeneous landscapes (e.g. for feeding, reproducing and overwintering; van Huizen 1977), and the importance of particular habitat combinations for a species’ survival at the landscape level remain unclear (Barbaro et al. 2007). Moreover, since many of the reported carabid responses to landscape features are species specific, more attention should be devoted to the individual species level, and not only for species of present conservation concern but also for common and widely distributed species, as well as to the ecological group level. Nonetheless, sampling strategies that avoid confounding effects are needed to clearly assess the respective weights of local and landscape factors and, at the landscape scale, the respective importance of composition and configuration on a species’ survival. Indeed, experimental landscapes (Davies and Margules 1998) or mensurative experiments (Hurlbert 1984) would be useful to disentangle these gradients that tend to be naturally correlated (Niemelä 2001; Fahrig 2003). Attention should also be paid to discrepancies between carabid species’ responses to landscape features across countries, possibly denoting that the impact of landscape structure on a particular species is likely to differ over its distribution range. Finally, in the current context of continuous landscape transformation, more emphasis should be given to the role of newly created habitats and abandoned areas across countries regarding carabid distribution, as well as to singular expanding elements and surrogate habitats, such as golf courses or private gardens in urban environments (Tanner and Gange 2005; Saarikivi et al. 2010), roads (Koivula and Vermeulen 2005; Melis et al. 2010; Yamada et al. 2010), power lines (Hollmen et al. 2008) and biomass crops (coppice with short and very-short rotation) in either open or forested landscapes. Eventually, potential mechanisms could be investigated by confronting the empirical data with models of dispersal and survival in heterogeneous landscapes (see e.g. Vermeulen and Opsteeg 1994; Pichancourt et al. 2006).
9 Concluding remarks
Carabids are among the most species-rich families of beetles, which has made them a natural focus of entomological research. Carabidologists are busy studying this evolutionarily successful group at several levels, from sub-cellular to supra-individual. Indeed, from the discovery of a pH receptor on the antennae of carabid beetles (Merivee et al. 2005; Milius et al. 2006) to cross-continental, landscape related research (Niemelä and Kotze 2009; Magura et al. 2010; Vanbergen et al. 2010) “carabidologists do it all”. They are helped by a reasonably solid taxonomy, even if evolutionary relationships are still undetermined.
Carabidology has contributed to several prominent ecological theories, including metapopulation theory (pioneering work by Piet den Boer and colleagues), and provides one of the best examples of a consistent, systematic study of the effects of urbanisation on biodiversity (Niemelä et al. 2002, and subsequent studies). These somewhat ad hoc examples are still powerful in the argumentation to encourage the use of carabids in ecological, evolutionary and behavioural studies.
Even from a subjective summary as this article admittedly is, it is obvious that carabids have contributed in a major way to our understanding of invertebrate adaptations, phylogeny and ecology. Accepting Hutchinson’s analogy that on the world stage an ecological play is being played out in the evolutionary theatre (Hutchinson 1965), watching and describing the peculiarities of one of the star players, ground beetles, will certainly advance our understanding of nature. In an age in which the earth is dominated by humans, this will provide important knowledge on how to maintain the richness of life on Earth, and with it, extend the lifespan of our own species.
Acknowledgements
We thank Thorsten Assmann and Piet den Boer for useful insights, Terry Erwin for information and reviewing this submission, and Ivailo Stoyanov and Lyubomir Penev for support and advice. For some authors, partial support was received from the Academy of Finland (JK, SV, project number 126915), the Czech Science Foundation (PS, grant # 206/09/1266), and the Ministerio de Ciencia e Innovación and the Fundación Séneca, Murcia (JS, respective project numbers: CGL2006–06706 and 08724PI08). The 14th ECM at Westerbork, the Netherlands received financial support from the Uyttenboogaart-Eliasen Foundation and the Province of Drente. The meeting was organised by volunteers of the Foundation Willem Beijerinck Biological Station (WBBS) and the SFOC – Dutch Carabidological Association (Loopkeverstichting). We especially thank Tim Opsteeg (WBBS) for designing and taking care of the website and Willem Jongbloed (WBBS) for taking care of the finances.
References
- Adis J, Paarmann W, Erwin TL. (1986) On the natural history and ecology of small terrestrial ground beetles (Col.: Tachyina: Polyderis) from an Amazonian blackwater inundation forest. In: Den Boer PJ, Luff ML, Mossakowski D, Weber F (Eds) Carabid beetles - Their Adaptations and Dynamics. Gustav Fischer Verlag, Stuttgart-New York, 413–427.
- Adis J, Paarmann W, Höfer H. (1990) On phenology and life cycle of Scarites (Scaritini, Carabidae, Coleoptera) from Central Amazonian floodplains. In: Stork NE (Ed) The Role of Ground Beetles in Ecological and Environmental Studies. Intercept, Andover, Hampshire, 269–275.
- Alexandrovitch OR, Lopatin IK, Pisanenko AD, Tsinkevitch VA, Snitko SM. (1996) A Catalogue of Coleoptera (Insecta) of Belarus. Minsk, FFR RB, 103 pp.
- Althoff G-H, Hockmann P, Klenner M, Nieheus F-J, Weber F. (1994) Dependence of running activity and net reproduction in Carabus auronitens on temperature. In: Desender K, Dufrêne M, Loreau M, Luff ML, Maelfait J-P (Eds) Carabid beetles: Ecology and EvolutionKluwer Academic Publishers, Dordrecht, The Netherlands, 95–100.
- Andersen J, Skorping A. (1990) Sympatric speciation by habitat specialization and parasitism in carabid beetles. In: Stork NE (Ed) The role of ground beetles in ecological and environmental studies. Intercept, Andover, 21–29.
- Anderson R, McFerran D, Cameron A. (2000) The ground beetles of Ireland (Coleoptera – Carabidae. Ulster Museum, 246 pp.
- Andorkó R, Kádár F. (2006) Carabid beetle (Coleoptera: Carabidae) communities in a woodland habitat in Hungary. Entomologica Fennica 17:221-228. [Google Scholar]
- Andrewartha HG, Birch LC. (1954) The distribution and abundance of animals. Chicago University Press, Chicago, 782 pp.
- Andújar A, Serrano J. (2001) Revisión y filogenia de los Zabrus Clairville, 1806 de la Península Ibérica (Colleoptera, Carabidae). Zaragoza, 90 pp.
- Angus RB, Galián J, Wrase DW, Chaladze G. (2008) The western Palaearctic species of the Pterostichus nigrita (Paykull) complex, with the description of a new species from Spain and a new subspecies of P. nigrita from Anatolia (Coleoptera, Carabidae). Nouvelle Revue d’Entomologie (N.S.) 25: 297–316. (published in 2009)
- Anonymous. (2006) Ground beetles of Ireland. National Museums Northern Ireland, Northern Ireland Environment Agency and the Heritage Council of Ireland. http://www.habitas.org.uk/groundbeetles/
- Antoine M. (1955–1962) Coléoptères Carabiques du Maroc. I-V. Mémoires de la Société de Sciences Naturelles et Physiques du Maroc, Zoologie (n.s.), Rabat, 694 pp.
- Apfelbeck V. (1904) Die Käferfauna der Balkanhabinsel 1 – Caraboidea. Friedländer, Berlin, 422 pp.
- Arndt E. (1991) Carabidae. In: Klausnitzer B: Die Larven der Käfer Mitteleuropas 1. Goecke & Evers, Krefeld, 45–141.
- Arndt E. (1998) Phylogenetic investigation of Carabidae (Coleoptera) using larval chaetotaxy. In: Ball GE, Casale A, Vigna Taglianti A (Eds) Phylogeny and classification of Caraboidea (Coleoptera: Adephaga). Museo Regionale di Scienze Naturali, Torino, 171–190.
- Arndt E. (2005) GROUND beetles (Coleoptera: Carabidae) as CROWN beetles in a Central European flood plain forest. In: Lövei GL, Toft S (Eds), European Carabidology 2003. Danish Institute of Agricultural Sciences 114:17-23. [Google Scholar]
- Arndt E, Kirmse S. (2002) Adaptation to seed-feeding in ground beetles (Coleoptera: Carabidae: Harpalini) of South Venezuela. Studies on Neotropical Fauna and Environment 37:151-157. doi: 10.1076/snfe.37.2.151.8581 [DOI] [Google Scholar]
- Arndt E, Paarmann W, Adis J. (1996) Description of larvae and larval specializations to a specific food in the genus Notiobia Perty (Coleoptera: Carabidae) from Amazonian lowlands. Studies on Neotropical Fauna and Environment 31:205-216. doi: 10.1076/snfe.31.3.205.13340 [DOI] [Google Scholar]
- Arndt E, Schnitter P, Sfenthourakis S, Wrase D. (in press) Groundbeetles (Carabidae) of Greece. Pensoft, Sofia, Moscow.
- Ås S. (1984) To fly or not to fly? Colonization of Baltic islands by winged and wingless carabid beetles. Journal of Biogeography 11:413-426. doi: 10.2307/2844805 [DOI] [Google Scholar]
- Ashworth AC. (1996) The response of arctic Carabidae (Coleoptera) to climate change based on the fossil record of the Quaternary Period. Annales Zoologici Fennici 33:125-131. [Google Scholar]
- Assmann T. (1990) Multiallelic genes in the carabid beetle Carabus punctatoauratus Germar from the Pyrenees. In: Stork NE (Ed) The role of ground beetles in ecological and environmental studies. Intercept, Andover, 319–324.
- Assmann T. (2009) Ground beetles and global change: first results from ongoing studies on case study species. In: Hengeveld R, Noordijk J, Opsteeg T, Turin H, Vermeulen HJW (Eds) Abstracts of the XIV European Carabidologists Meeting, Westerbork, the Netherlands, 25.
- Assmann T, Buse J, Drees C, Habel J, Härdtle W, Matern A, Von Oheimb G, Schuldt A, Wrase DW. (2008) From Latreille to DNA systematics – towards a modern synthesis for carabidology. In: Penev L, Erwin TL, Assmann T (Eds) Back to the roots and back to the future. Pensoft, Sofia, 41–76.
- Assmann T, Dormann W, Främbs H, Gürlich S, Handke K, Huk T, Terlutter H. (2002) Rote liste der Niedersachsen und Bremen gefährdeten Sandlaufkäfer und Laufkäfer (Coleoptera: Cicindelidae et Carabidae) mit Gesamtartenverzeichnis. 1. Fassung vom 1.6.2002. Niedersächsiches Landesamt für Ökologie.
- Assmann T, Nolte O, Reuter H. (1994) Postglacial colonization of middle Europe by Carabus auronitens, as revealed by population genetics (Coleoptera, Carabidae). In: Desender K, Dufrêne M, Loreau M, Luff ML, Maelfait J-P (Eds) Carabid beetles: Ecology and evolution. Kluwer Academic Press, Dordrecht, 3–9.
- Aukema B. (1990) Wing length determination in two wing-dimorphic Calathus species (Coleoptera, Carabidae). Hereditas113: 189–202. doi: 10.1111/j.1601-5223.1990.tb00084.x [DOI]
- Aukema B. (1991) Fecundity in relation to wing-morph of three closely related species of the melanocephalus group of the genus Calathus (Coleoptera, Carabidae). Oecologia 87:118-126. doi: 10.1007/BF00323789 [DOI] [PubMed] [Google Scholar]
- Aukema B. (1995) Flying for life: Wing dimorphism in closely related species of the genus Calathus (Coleoptera: Carabidae) Ph.D. thesis, Wageningen University, Ponsen & Loojien, Wageningen, The Netherlands, 168 pp.
- Aviron S, Burel F, Baudry J, Schermann N. (2005) Carabid assemblages in agricultural landscapes: impacts of habitat features, landscape context at different spatial scales and farming intensity. Agriculture, Ecosystems and Environment 108:205-217. doi: 10.1016/j.agee.2005.02.004 [DOI] [Google Scholar]
- Baars MA. (1982) Running for life. Thesis, University of Amsterdam, 161 pp.
- Baars MA, Van Dijk ThS. (1984) Population dynamics of two carabid beetles at a Dutch heathland. II. Egg production and survival in relation to density. Journal of Animal Ecology 53:389-400. doi: 10.2307/4523 [DOI] [Google Scholar]
- Baehr M. (1994) Phylogenetic relations and biogeography of the genera of Pseudomorphinae (Coleoptera, Carabidae). In: Desender K, Dufrêne M, Loreau M, Luff ML, Maelfait J-P (Eds) Carabid beetles: Ecology and evolution. Kluwer Academic Press, Dordrecht, 11–17.
- Baehr M. (1998) A preliminary survey of the classification of the Psydrinae (Coleoptera: Carabidae). In: Ball GE, Casale A, Vigna Taglianti A (Eds) Phylogeny and classification of Caraboidea (Coleoptera: Adephaga). Museo Regionale di Scienze Naturali, Torino, 359–367.
- Ball GE. (1979) Conspectus of carabid classification: history, holomorphology, and higher taxa. In: Erwin TL, Ball GE, Whitehead DR, Halpern AL (Eds) Carabid Beetles: Their Evolution, Natural History, and Classification. Dr. W. Junk Publishers, Dordrecht, Boston, Lancaster, 63–111.
- Ball GE (Ed) (1985) Taxonomy, phylogeny and zoogeography of beetles and ants: a volume dedicated to the memory of Philip Jackson Darlington, Jr (1904–1983). Dr. W. Junk Publishers, Dordrecht, Boston, Lancaster: 513 pp.
- Ball GE, Bousquet Y. (2001) Carabidae. In: Arnett RH, Thomas RC (Eds) American Beetles, Volume I: Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC Press LLC, Boca Raton, FL, 32–132.
- Ball GE, Casale A, Vigna Taglianti A.Eds (1998) Phylogeny and classification of Caraboidea (Coleoptera: Adephaga). Museo Regionale di Scienze Naturali, Torino, 543 pp.
- Bangsholt F. (1983) The distribution and occurrence of tiger beetles and ground beetles in Denmark from about 1830 to 1981 (Coleoptera: Cicindelidae and Carabidae). Scandinavian Science Press, Copenhagen, pp. 271.
- Barbaro L, Pontcharraud L, Vetillard F, Guyon D, Jactel H. (2005) Comparative responses of bird, carabid, and spider assemblages to stand and landscape diversity in maritime pine plantation forests. Ecoscience 12:110-121. doi: 10.2980/i1195-6860-12-1-110.1 [DOI] [Google Scholar]
- Barbaro L, Rossi J-P, Vetillard F, Nezan J, Jactel H. (2007) The spatial distribution of birds and carabid beetles in pine plantation forests: the role of landscape composition and structure. Journal of Biogeography 34:652-664. doi: 10.1111/j.1365-2699.2006.01656.x [DOI] [Google Scholar]
- Barbaro L, van Halder I. (2009) Linking bird, carabid beetle and butterfly life-history traits to habitat fragmentation in mosaic landscapes. Ecography 32:321-333. doi: 10.1111/j.1600-0587.2008.05546.x [DOI] [Google Scholar]
- Barber H. (1931) Traps for cave-inhabiting insects. Journal of the Elisha Mitchell Scientific Society 46:259-266. [Google Scholar]
- Barsevskis A. (2003) Latvijas Skrejvaboles (Coleoptera: Carabidae, Trachypachidae & Rhysodidae). Baltic Institute of Coleopterology, Daugavpils, 262 pp.
- Barton PS, Manning AD, Gibb H, Lindenmayer DB, Cunnigham SA. (2009) Conserving ground-dwelling beetles in an endangered woodland community: Multi-scale habitat effects on assemblage diversity. Biological Conservation 142:1701-1709. doi: 10.1016/j.biocon.2009.03.005 [DOI] [Google Scholar]
- Bauer T, Talarico F, Mazzei A, Giglio A, Zetto Brandmayr T, Brandmayr P, Betz O. (2005) Hunting ants in Mediterranean clay soils: life history of Siagona europaea (Coleoptera, Carabidae). Italian Journal of Zoology 72:33-42. doi: 10.1080/11250000509356650 [DOI] [Google Scholar]
- Bedel L. (1899–1900) Carabidae. In: Catalogue raisonné des Coléoptères du Nord de l’Afrique. Publication de la Société entomologique de France, Paris, I-II: 1–320.
- Bell RT. (1998) Where do the Rhysodini (Coleoptera) belong? In: Ball GE, Casale A, Vigna Taglianti A (Eds) Phylogeny and classification of Caraboidea (Coleoptera: Adephaga). Museo Regionale di Scienze Naturali, Torino, 261–271.
- Bérces S, Szél G, Ködöböcz V, Kutasi C. (2008) The distribution, habitat, and the nature conservation value of a Natura 2000 beetle, Carabus hungaricus Fabricius, 1972 in Hungary. In: Penev L, Erwin T, Assmann T (Eds) Back to the roots and back to the future. Towards a new synthesis amongst taxonomic, ecological and biogeographical approaches in carabidology. Proceedings of the XIII European Carabidologists Meeting, Blagoevgrad, August 20–24, 2007, 363–372.
- Beutel R. (1998) Trachypachidae and the phylogeny of Adephaga (Coleoptera). In: Ball GE, Casale A, Vigna Taglianti A (Eds) Phylogeny and classification of Caraboidea (Coleoptera: Adephaga). Museo Regionale di Scienze Naturali, Torino, 81–105.
- Beutel RG, Haas A. (1996) Phylogenetic analysis of larval and adult characters of Adephaga (Coleoptera) using cladistic computer programs. Entomologica Scandinavica 27:197-205. [Google Scholar]
- Blake S, Foster GN, Fisher EJ, Ligertwood GL. (1996) Effects of management practices on the carabid faunas of newly established wildflower meadows in southern Scotland. Annales Zoologici Fennici 33:139-147. [Google Scholar]
- Bodenheimer FS. (1934) Studies on the ecology of Palestinian Coleoptera. II. Seasonal and diurnal appearance and activity. Bulletin de la Société Royale Entomologique d’Egypt 18:211-241. [Google Scholar]
- Boeken M, Desender K, Drost B, Van Gijzen T, Koese B, Muilwijk J, Turin H, Vermeulen R. (2002) De loopkevers van Nederland en Vlaanderen (Coleoptera, Carabidae). Jeugdbondsuitgeverij, Nederland, 212 pp.
- Bommarco R. (1998) Reproduction and energy reserves of a predatory carabid beetle relative to agroecosystem complexity. Ecological Applications 8:846-853. doi: 10.1890/1051-0761(1998)008[0846:RAEROA]2.0.CO;2 [DOI] [Google Scholar]
- Bonavita P, Chemini C. (1996) Structures and indicator role of carabid assemblages from wet areas in the province of Trento, Italian Alps (Coleoptera, Carabidae). Quaderni della Stazioni diEcologia del civico Museo di Storia naturale di Ferrera 10:107-123. [Google Scholar]
- Borcherding R, Paarmann W, Bin Nyawa S, Bolte H. (2000) How to be a fig beetle? Observations of ground beetles (Col., Carabidae) associated with fruitfalls in a rain forest of Borneo. Ecotropica 6:169-180. [Google Scholar]
- Bouget C. (2004) Chablis et diversité des coléoptères en forêt feuillue de plaine : impact à court terme de la trouée, de sa surface et de son contexte paysager. PhD thesis in Ecology, Paris, France, Muséum National d’Histoire Naturelle, 452 pp.
- Bouget C. (2005) Short-term effect of windthrow disturbance on ground beetle communities: gap and gap size effects. In: Lövei G, Toft S (Eds), European Carabidology 2003, 11th European Carabidologists' Meeting. Aarhus (Denmark), July 2003, Danish Institute of Agricultural Sciences, DIAS Report 114, 25–40.
- Bouget C, Duelli P. (2004) The effects of windthrow on forest insect communities: a literature review. Biological Conservation 118:281-299. doi: 10.1016/j.biocon.2003.09.009 [DOI] [Google Scholar]
- Bowie MH, Frampton CM. (2004) A practical technique for non-destructive monitoring of soil surface invertebrates for ecological restoration programmes. Ecological Management & Restoration 5:34-42. doi: 10.1111/j.1442-8903.2004.00171.x [DOI] [Google Scholar]
- Brakman PJ. (1966) Lijst van Coleoptera uit Nederland en het omliggend gebied. Monografieën van de Nederlandse Entomologische Vereniging 2, Amsterdam, 219 pp.
- Brandmayr P. (1977) Ricerche etologiche e morfofunzionali sulle cure parentali in Carabidi Pterostichini (Coleoptera, Carabidae, Pterostichinae). Redia 60:275-316. [Google Scholar]
- Brandmayr P, Den Boer PJ, Weber F.Eds (1983) The synthesis of field study and laboratory experiment. Proceedings of the 4th European Carabidologists’ Meeting. Centre for Agricultural Publishing and Documentation Wageningen, the Netherlands, 196 pp.
- Brandmayr P, Lövei GL, Zetto Brandmayr T, Casale A, Vigna Taglianti A.Eds (2000) Natural history and applied ecology of carabid beetles. Proceedings of the 9th European Carabidologists’ Meeting. Pensoft, Sofia-Moscow, 304 pp.
- Brandmayr P, Zetto Brandmayr T. (1974) Sulle cure parentali e su altri aspetti della biologia di Carterus (Sabienus) calydonius Rossi, con alcune considerazioni sui fenomeni di cura della prole sino ad oggi riscontrati in Carabidi (Coleoptera, Carabidae): Redia 55: 143–175.
- Brandmayr P, Zetto Brandmayr T. (1979) The evolution of parental care phenomena in Pterostichini, with particular reference to the genera Abax and Molops. In: Den Boer PJ, Thiele HU, Weber F (Eds) On the evolution of behaviour in carabid beetles. Miscellaneous Papers, 18, Landbouwhogeschool Wageningen, 35–49.
- Brandmayr P, Zetto Brandmayr T. (1982) Identificazione di larve del genere Ophonus Dejean, 1821 (sensu novo) e note bionomiche. Memorie di Società entomologica Italiana 60 (1981):67-103. [Google Scholar]
- Brandmayr P, Zetto Brandmayr T. (1987) The problem of presocial behaviour in ditomine ground beetles. Pubblicazioni dell'Istituto di Entomologia dell'Universita di Pavia 36:15-18. [Google Scholar]
- Brandmayr P, Zetto Brandmayr T. (1991) A paleoclimatic model for the evolution of brood watching in the ground beetle genus Molops (Coleoptera, Carabidae). Ethology Ecology & Evolution, Special issue 1:139-142. [Google Scholar]
- Brandmayr P, Zetto Brandmayr T. (1994) The evolutionary history of the genus Abax (Coleoptera, Carabidae). In: Desender K, Dufrêne M, Loreau M, Luff ML, Maelfait J-P (Eds) Carabid beetles: ecology and evolution. Kluwer Academic Press, Dordrecht, 19–24.
- Braun SD, Jones TH, Perner J. (2004) Shifting average body size during regeneration after pollution - a case study using ground beetle assemblages. Ecological Entomology 29:543-554. doi: 10.1111/j.0307-6946.2004.00643.x [DOI] [Google Scholar]
- Bräuniger C, Knapp S, Kuhn I, Klotz S. (2010) Testing taxonomic and landscape surrogates for biodiversity in an urban setting. Landscape and Urban Planning 97:283-295. doi: 10.1016/j.landurbplan.2010.07.001 [DOI] [Google Scholar]
- Breuning S. (1932–1937) Monographie der Gattung Carabus, I-VII. Bestimmungstabellen der Europäischen Käfer 104, 105 (1932) 106, 107 (1933) 108 (1934) 109 (1935) 110 (1937). Troppau 1–1610 (+ 41 distribution maps).
- Brose U. (2003) Island biogeography of temporary wetland carabid beetle communities. Journal of Biogeography 30:879-888. doi: 10.1046/j.1365-2699.2003.00893.x [DOI] [Google Scholar]
- Brouat C, Sennedot F, Audiot P, Leblois R, Rasplus J-Y. (2003) Fine-scale genetic structure of two carabid species with contrasted levels of habitat specialization. Molecular Ecology 12:1731-1745. doi: 10.1046/j.1365-294X.2003.01861.x [DOI] [PubMed] [Google Scholar]
- Brückner M, Mossakowski D. (2006) Phylogeny of the genus Percus (Coleoptera: Carabidae) - nuclear genes and the basal splits. Entomologica Fennica 17:195-199. [Google Scholar]
- Brunsting AMH, Siepel H, Van Schaik Zillesen PG. (1986) The role of larvae in the population ecology of Carabidae. In Den Boer PJ, Luff ML, Mossakowski D, Weber F (Eds) Carabids Beetles, Their Adaptations and Dynamics. Gustav Fischer, Stuttgart New York, 399–411.
- Brust GE, House GJ. (1988) Weed seed destruction by arthropods and rodents in low-input soybean agroecosystems. American Journal of Alternative Agriculture 3:19-25. doi: 10.1017/S0889189300002083 [DOI] [Google Scholar]
- Burakowski B, Mroczkowski M, Stevanska J. (1973–1974) Carabidae 1–2. Katalog Fauny Polski, 23: 1–232, 1430.
- Burel F. (1989) Landscape structure effects on carabid beetles spatial patterns in Western France. Landscape Ecology 2:215-226. doi: 10.1007/BF00125092 [DOI] [Google Scholar]
- Burel F, Baudry J, Butet A, Clergeau P, Delettre Y, Le Coeur D, Dubs F, Morvan N, Paillat G, Petit S, Thenail C, Brunel E, Lefeuvre JC. (1998) Comparative biodiversity along a gradient of agricultural landscapes. Acta Oecologica 19:47-60. doi: 10.1016/S1146-609X(98)80007-6 [DOI] [Google Scholar]
- Burel F, Butet A, Delettre YR, Millàn de la Peña N. (2004) Differential response of selected taxa to landscape context and agricultural intensification. Landscape and Urban Planning 67:195-204. doi: 10.1016/S0169-2046(03)00039-2 [DOI] [Google Scholar]
- Butterfield J. (1997) Carabid community succession during the forestry cycle in conifer plantations. Ecography 20:614-625. doi: 10.1111/j.1600-0587.1997.tb00430.x [DOI] [Google Scholar]
- Cabidoche M. (1963) Note sur la périodicité saissonière d’activité d’une poupulation de Coléoptères troglobies (Aphaenops et Hydraphaenops) Comptes Rendus de Académie des Sciences Paris, 256: 4491–4493.
- Cabidoche M. (1966) Contribution à la connaissance de l’écologie des Trechinae cavernicoles pyrénèens. PhD thesis, University Paris, 228 pp.
- Callot HJ, Schott C. (1993) Catalogue et atlas des coleopteres d’Alsace. Societé Alsacienne d’Entomologie, 122 pp.
- Casale A. (1988) Revisione degli Sphodrina (Coleoptera, Carabidae, Sphodrini). Museo Regionale di Scienze naturali, Torino, 1024 pp.
- Casale A. (1998) Phylogeny and biogeography of Calleidina (Coleoptera: Carabidae: Lebiini): a preliminary survey. In: Ball GE, Casale A, Vigna Taglianti A (Eds) Phylogeny and classification of Caraboidea (Coleoptera: Adephaga). Museo Regionale di Scienze Naturali, Torino, 381–427.
- Casale A. (2005) Tabelle per l’identificazione delle tribu e dei Generi di Caraboidea presenti in Italia. In: Brandmayr P, Zetto T, Pizzolotto, R (Eds) I Colleotteri Carabidi per la valutazione ambientale e la conservazione della biodiversità. APAT, Roma, 148–185.
- Casale A, Busato E. (2008) A real time extinction: the case of Carabus clatratus in Italy (Coleoptera, Carabidae). In: Penev L, Erwin TL, Assmann T (Eds) Back to the roots and back to the future. Pensoft, Sofia, 353–362.
- Casale A, Mossakowski D. (2003) General remarks on the classification of Imura Y (2002). In: Turin H, Penev L, Casale A (Eds) The genus Carabus - A Synthesis. Sofia-Moscow: 455–461.
- Casale A, Prüser E, Arndt E, Mossakowski D. (1998) Phylogenetic relationships in the subgenus Platycarabus Morawitz, 1886 (Coleoptera: Carabidae: Carabini). In: Ball GE, Casale A, Vigna Taglianti A (Eds) Phylogeny and classification of Caraboidea (Coleoptera: Adephaga). Museo Regionale di Scienze Naturali, Torino, 429–447.
- Casale A, Sturani M, Vigna Taglianti A. (1982) Fauna d’Italia – Carabidae I – Introduzione, Paussinae, Carabinae. Edizione Calderini, Bologna, 499 pp.
- Casale A, Vigna Taglianti A, Brandmayr P, Colombetta G. (2007) Insecta Coleoptera Carabidae (Carabini, Cychrini, Trechini, Abacetini, Stomini, Pterostichini). In: Ruffo S, Stoch F (Eds) Checklist and distribution of the Italian Fauna. Memorie del Museo Civico di Storia Naturale di Verona 2 serie, Sezione Scienze della Vita 17 (2006): 159–163 with data on CD-ROM.
- Cavazzuti P. (1989) Monografia del Genere Procerus (Coleoptera, Carabidae, Carabini). Associazione Naturalistica Piemontese, Memorie Vol. I. Edizione L’Artistica Savigliano, 200 pp.
- Charrier S, Petit S, Burel F. (1997) Movements of Abax parallelepipedus (Coleoptera, Carabidae) in woody habitats of a hedgerow network landscape: a radio-tracing study. Agriculture, Ecosystems and Environment 61:133-144. doi: 10.1016/S0167-8809(96)01101-2 [DOI] [Google Scholar]
- Cole LJ, Pollock ML, Robertson D, Holland JP, McCracken DI. (2006) Carabid (Coleoptera) assemblages in the Scottish uplands: the influence of sheep grazing on ecological structure. Entomologica Fennica 17:229-240. [Google Scholar]
- Coulon J, Marchal P, Pupier R, Richoux P, Allemand R, Genest L-C, Clary J. (2000) Coléoptères de Rhône-Alpes. Société linnéenne de Lyon, 193 pp.
- Crawley MJ. (2003) Statistical Computing. An Introduction to Data Analysis using S-Plus. John Wiley & Sons Ltd., England, 761 pp.
- Csiki E. (1946) Die Käferfauna des Karpaten-Beckens 1. Algemeiner Teil und Caraboidea. Naturwissenschaftliche Monographien IV. András Tasnádi Kubacska, Budapest, 798 pp.
- Cuesta D, Taboada A, Calvo L, Salgado JM. (2006) A preliminary investigation of ground beetle (Coleoptera: Carabidae) assemblages and vegetation community structure in Calluna vulgaris heathlands in NW Spain. Entomologica Fennica 17:241-252. [Google Scholar]
- Czechowski W. (1982) Occurrence of carabids Coleoptera, Carabidaein the urban greenery of Warsaw according to the land utilization and cultivation. Memorabilia Zoologica39: 3–108.
- Dalthorp D. (2004) The generalized linear model for spatial data: assessing the effects of environmental covariates on population density in the field. Entomologia Experimentalis et Applicata 111:117-131. doi: 10.1111/j.0013-8703.2004.00158.x [DOI] [Google Scholar]
- Darlington PJ. (1936) Variation and atrophy of flying wings of some carabid beetles Annals of the Entomological Society of America49: 136–179.
- Darlington PJ. (1943) Carabidae of mountains and islands: data on the evolution of isolated faunas, and on atrophy of wings. Ecological Monographs13: 37–61. doi: 10.2307/1943589 [DOI]
- Davies K, Margules C. (1998) Effects of habitat fragmentation on carabid beetles: Experimental evidence. Journal of Animal Ecology 67:460-471. doi: 10.1046/j.1365-2656.1998.00210.x [DOI] [Google Scholar]
- Davies L. (1972) Two Amblystogenium species (Col., Carabidae) coexisting on the Subarctic Possesion Island; Crozet Islands. Entomologica Scandinavica 3:275-286. [Google Scholar]
- Darwin C. (1859) On the origin of species by means of natural selection or the preservation of favoured races in the struggle for life. Hazel, Watson & Viney, London, 389 pp. [PMC free article] [PubMed]
- Den Boer PJ. (1968) Spreading of risk and stabilization of animal numbers. Acta Biotheoretica 18:165-194. doi: 10.1007/BF01556726 [DOI] [PubMed] [Google Scholar]
- Den Boer PJ. (1970) On the significance of dispersal power for populations of carabid-beetles (Coleoptera, Carabidae). Oecologia 4:1-28. doi: 10.1007/BF00390612 [DOI] [PubMed] [Google Scholar]
- Den Boer PJ. (Ed)(1971) Dispersal and dispersal power of carabid beetles Proceedings of the 1st European Carabidologists’ Meeting. Miscellaneous papers, Agricultural University Wageningen 8: 151 pp.
- Den Boer PJ. (1977) Dispersal power and survival. Carabids in a cultivated countryside. Miscellaneous papers, Agricultural University, Wageningen 14:1-190. [Google Scholar]
- Den Boer PJ. (1980) Exclusion or coexistence and the taxonomic or ecological relationship between species. Netherlands Journal of Zoology 30:278-306. doi: 10.1163/002829679X00421 [DOI] [Google Scholar]
- Den Boer PJ. (1985) Exclusion, competition or co-existence? A question for testing the right hypothesis. Zeischrift für zoologische Systematik und Evolution 23:259-274. [Google Scholar]
- Den Boer PJ. (1986) Facts, hypotheses and models on the part played by food in the dynamics of carabid populations. In Den Boer PJ, Grüm L, Szyszko J (Eds) Feeding behaviour and accessibility of food for carabid beetles. Proceedings of the 5th Meeting of European Carabidologists’ Meeting. Warsaw Agricultural University Press, Warsaw, 81–96.
- Den Boer PJ, Den Boer-Daanje W. (1990) On life history tactics in carabid beetles: are there only spring and autumn breeders. In: Stork NE (Ed) The Role of Ground Beetles in Ecological and Environmental Studies, 247–258.
- Den Boer PJ, Gradwell GR.Eds (1971) Dynamics of populations, proceedings of the advanced study institute on Dynamics of Numbers in Populations, Oosterbeek, the Netherlands 7–18 September 1970, Centre for agricultural publishing and documentation, Wageningen, 611 pp.
- Den Boer PJ, Grüm L, Szyszko J.Eds (1986a) Feeding behaviour and accessibility of food for carabid beetles. Proceedings of the 5th European Carabidologists’ Meeting. Warsaw Agricultural University Press, Warsaw, 167 pp.
- Den Boer PJ, Luff ML, Mossakowski D, Weber F.Eds (1986b) Carabid Beetles, their adaptations and dynamics. Proceedings of the 17th International Entomological Congress. Fischer, Stuttgart-New York, 551 pp.
- Den Boer PJ, Lövei GL, Stork NE, Sunderland KD.Eds (1987) Proceedings of the 6th European Carabidologist’s Meeting. Hungary. Phytopthologica et Entomologica Hungarica 22:1-458. [Google Scholar]
- Den Boer PJ, Mols PJM, Szyszko J.Eds (1993) Dynamics of populations, proceedings of the meeting on population problems, Smolarnia, Poland 9–15 September 1992. Warsaw Agricultural University Press, Warsaw, 111 pp.
- Den Boer PJ, Reddingius J. (1996) Regulation and stabilization paradigms in population ecology, population and community biology series 16: 1–397, Chapman & Hall, London.
- Den Boer PJ, Thiele H-U, Weber F.Eds (1979) On the evolution of behaviour in Carabid beetles. Proceedings of the 3rd European Carabidologists’ Meeting. Miscellaneous papers, Agricultural University Wageningen 18: 222 pp.
- Den Boer PJ, Van Dijk ThS. (1994) Carabid beetles in a changing environment. Wageningen Agricultural University Papers, Wageningen, the Netherlands, 94:1-30. [Google Scholar]
- Dennis RLH, Shreeve TG, Sheppard DA. (2007) Species conservation and landscape management: a habitat perspective. In: Stewart AJA, New TR, Lewis OT (Eds) Insect conservation biology, CABI, Wallingford, 92–126. doi: 10.1079/9781845932541.0092 [DOI]
- Desender K. (1986) Distribution and ecology of carabid beetles in Belgium (Coleoptera, Carabidae), Parts 1–4. Institut Royal des Sciences Naturelles de Belgique. Documents de Travail 26: 1–30, 27: 1–24, 30: 1–23, 34:1-48. [Google Scholar]
- Desender K. (1989a) Heritability of wing development and body size in a carabid beetle, Pogonus chalceus Marsham, and its evolutionary significance. Oecologia 78:513-520. doi: 10.1007/BF00378743 [DOI] [PubMed] [Google Scholar]
- Desender K. (1989b) Dispersievermogen en ecologie van loopkevers (Coleoptera, Carabidae) in Belgie: een evolutionaire benadering. Royal Belgian Institute of Natural Sciences and Royal Belgian Institute of Natural Sciences, Brussels, Belgium, 136 pp.
- Desender K. (2000) Flight muscle development and dispersal in the life cycle of carabid beetles: patterns and processes Bulletin de l’Institut Royal des Sciences Naturelles de Belgique, Entomologie 70: 13–31.
- Desender K. (2005) Theory versus reality: a review on the ecological and population genetic effects of forest fragmentation on wild organisms, with an emphasis on ground beetles. DIAS report 114:49-72. [Google Scholar]
- Desender K, Baert L, Maelfait J-P. (1990) Evolutionary ecology of carabids in the Galapagos Archipelago. In: Stork NE (Ed) The role of ground beetles in ecological and environmental studies. Intercept, Andover, 13–20.
- Desender K, Dekoninck W, Dufrêne M, Maes D. (2010) Changes in the distribution of carabid beetles in Belgium revisited: have we halted the diversity loss? Biological Conservation 143: 1549–1557. doi: 10.1016/j.biocon.2010.03.039 [DOI]
- Desender K, Dekoninck W, Maes D, Crevecoeur L, Dufrêne M, Jacobs M, Lambrechts J, Pollet M, Stassen E, Thys N. (2008a) Een nieuwe verspreidingsatlas van de loopkevers en zandloopkevers (Carabidae) in Belgie. Rapporten van het Instituut voor Natuur- en Bosonderzoek 2008 (INBO.R.2008.13), Instituut voor Natuur- en Bosonderzoek, Brussels, 184 pp.
- Desender K, Dekoninck W, Maes D. (2008b) An updated Red List of the ground and tiger beetles (Coleoptera, Carabidae) in Flanders. Bulletin De L'Institut Royal Des Sciences Naturelles De Belgique, Entomologie 78:113-131. [Google Scholar]
- Desender K, Dufrêne M, Loreau M, Luff ML, Maelfait JP.Eds (1994a) Carabid Beetles: Ecology and Evolution, Proceedings of the 8th European Carabidologists’ Meeting. Kluwer Academic Publishers, Dordrecht, the Netherlands, 474 pp.
- Desender K, Dufrêne M, Maelfait JP. (1994b) Long-term dynamics of carabid beetles in Belgium: a preliminary analysis on the influence of changing climate and land use by means of a database covering more than a century. In: Desender K, Dufrêne M, Loreau M, Luff ML, Maelfait JP (Eds) Carabid Beetles: Ecology and Evolution, Kluwer Academic Publishers, Dordrecht, the Netherlands, 247–252.
- Desender K, Maes D, Maelfait J-P, Van Kerckvoorde M. (1995) Een gedocumenteerde Rode Lijst van de zandloopkever en loopkevers van Vlaanderen. Instituut voor Natuurbeboud, Hasselt, 208 pp.
- Desender KRC, Serrano J, Verdyck P. (2000) Genetic diversity and wing polymorphism in the saltmarsh beetle Pogonus chalceus: an Atlantic-Mediterranean comparison. In: Brandmayr P, Lövei GL, Zetto Brandmayr T, Casale A, Vigna Taglianti A (Eds) Natural history and applied ecology of carabid beetles. Pensoft, Sofia, 35–43.
- Desender K, Small E, Gaublomme E, Verdyck P. (2005) Rural-urban gradients and the population genetic structure of woodland ground beetles. Conservation Genetics 6:51-62. doi: 10.1007/s10592-004-7748-3 [DOI] [Google Scholar]
- Desender K, Turin H. (1989) Loss of habitat and changes in the composition of the ground and tiger beetle fauna in four West European countries since 1950 (Coleoptera: Carabidae, Cicindelidae). Biological Conservation 48:277-294. doi: 10.1016/0006-3207(89)90103-1 [DOI] [Google Scholar]
- Desender K, Verdyck P. (2000) Genetic differentiation in the Galápagos caterpillar hunter Calosoma granatense (Coleoptera, Carabidae). In: Brandmayr P, Lövei G, Zetto Brandmayr T, Casale A, Vigna Taglianti A (Eds) Natural History and Applied Ecology of Carabid Beetles, Pensoft Publishers, Sofia, Bulgaria, 25–34.
- Deuve T. (1993) L’abdomen et les genitalia des femelles de Coléoptères Adephaga. Mémoires du Muséum National d’Histoire Naturelle 155: 184.
- Deuve T. (2004) Illustrated catalogue of the Genus Carabus of the world (Coleoptera: Carabidae). Pensoft, Sofia-Moscow, 461 pp.
- De Vries HH. (1994) Size of habitat and presence of ground beetle species In: Desender K, Dufrêne M, Loreau M, Luff ML, Maelfait J-P (Eds) Carabid beetles: ecology and evolution. Series Entomologica 51:253-259. [Google Scholar]
- De Vries HH, Den Boer PJ and Van Dijk TS. (1996) Ground beetle species in heathland fragments in relation to survival, dispersal, and habitat preference. Oecologia 107:332-342. doi: 10.1007/BF00328449 [DOI] [PubMed] [Google Scholar]
- Diaz M. (1994) Variability in seed size selection by granivorous passerines - effects of bird size, bird size variability, and ecological plasticity. Oecologia 99:1-6. doi: 10.1007/BF00317076 [DOI] [PubMed] [Google Scholar]
- Dieckmann U, O’Hara B, Weisser W. (1999) The evolutionary ecology of dispersal. Trends in Ecology and Evolution 14:88-90. doi: 10.1016/S0169-5347(98)01571-7 [DOI] [Google Scholar]
- Di Giulio A, Moore W. (2004) The first-instar larva of the genus Arthropterus (Coleoptera: Carabidae: Paussinae): implications for evolution of myrmecophily and phylogenetic relationships within the subfamily. Invertebrate Systematics 18:101-115. doi: 10.1071/IS03028 [DOI] [Google Scholar]
- Di Giulio A, Vigna Taglianti A. (2001) Biological observations on Pachyteles larvae (Coleoptera Carabidae Paussinae). Tropical Zoology 14:157-173. [Google Scholar]
- Digweed SC, Currie CR, Carcamo HA, Spence JR. (1995) Digging out the “digging-in effect” of pitfall traps: Influences depletion and disturbance on catches of ground beetles (Coleoptera: Carabidae). Pedobiologia 39:561-576. [Google Scholar]
- Dinter K, Paarmann W, Peschke K, Arndt E. (2002) Ecological, behavioural and chemical adaptations to ant predation in species of Thermophilum and Graphipterus (Coleoptera: Carabidae) in the Sahara desert. Journal of Arid Environment 50:267-286. doi: 10.1006/jare.2001.0850 [DOI] [Google Scholar]
- Drees C, Hüfner S, Matern A, Nève G, Assmann T. (2011)Repeated sampling detects gene flow in a flightless ground beetle in a fragmented landscape. Oikos accepted. [DOI] [PubMed]
- Drees C, Matern A, Vermeulen R, Assmann T. (2007) The influence of habitat quality on populations: a plea for an amended approach in the conservation of Agonum ericeti. Baltic Journal of Coleopterology 7:1-8. [Google Scholar]
- Drovenik B. (1999) Catalogus Faunae – Carabiden der Balkanländer, Coleoptera, Carabidae. Schwanfelder coleopterologische Mitteilungen 15:1-123. [Google Scholar]
- Du Bus de Warnaffe G, Lebrun P. (2004) Effects of forest management on carabid beetles in Belgium: implications for biodiversity conservation. Biological Conservation 118:219-234. doi: 10.1016/j.biocon.2003.08.015 [DOI] [Google Scholar]
- Du Chatenet G. (1986) Guide des Coléoptères d’Europe – Carabidae. Delachaux & Niestlé, Neuchâtel – Paris: 60–189, 28 plates, 190 maps.
- Duelli P, Obrist MK. (2003) Biodiversity indicators: the choice of values and measures. Agriculture, Ecosystems and Environment 98:87-98. doi: 10.1016/S0167-8809(03)00072-0 [DOI] [Google Scholar]
- Dufrêne M, Legendre P. (1997) Species assemblages and indicator species: the need for a flexible assymetrical approach. Ecological Monographs 67:345-366. [Google Scholar]
- Düring A. (2002) Differentiation between Abax parallelepipedus and Abax angustatus in morphological and genetic character complex. In: Szyszko J, Den Boer PJ, Bauer T (Eds) How to protect or what we know about carabid beetles. Warsaw Agricultural University Press, Warsaw, 37–44.
- Düring A. (2004) Molekularsystematische Untersuchungen am Carabidentaxob Abax Bonelli 1810 (Coleoptera, Carabidae), Eine phylogenetische und phylogeographische Studie. Verlagsgruppe Mainz GmbH, Aachen, 363 pp.
- Düring A, Brückner M. (2000) The evolutionary history of the tribe Molopini: a.first molecular approach. In: Brandmayr P, Lövei GL, Zetto Brandmayr T, Casale A, Vigna Taglianti A (Eds) Natural history and applied ecology of carabid beetles. Pensoft, Sofia, 1–4.
- Düring A, Brückner M, Mossakowski D. (2006) Different behaviour of mitochondrial and nuclear markers: introgression and the evolutionary history of Chrysocarabus (Coleoptera: Carabidae). Entomologica Fennica 17:200-206. [Google Scholar]
- Düring A, Bruckner M, Zimmermann M, Bobenhausen I, Eicke N, Mossakowski D. (2000) Geographic distribution of two highly different mtDNA haplotypes in Carabus (Chrysocarabus) splendens Olivier. In: Brandmayr P, Lövei GL, Zetto Brandmayr T, Casale A, Vigna Taglianti A (Eds) Natural history and applied ecology of carabid beetles. Pensoft, Sofia, 5–9.
- Dymitryszyn I, Szyszko J, Gryuntal S. (2003) Carabids (Carabidae, Col.) in the area of the planned A-2 highway section Poznan-Wrzesnia. Annals of Warsaw Agricultural University - SGGW, Horticulture and Landscape Architecture 24:149-154. [Google Scholar]
- Eggers B, Matern A, Drees C, Eggers J, Härdtle W, Assmann T. (2010) Value of semi-open corridors for simultaneously connecting open and wooded habitats: a case study with ground beetles. Conservation Biology 24:256-266. doi: 10.1111/j.1523-1739.2009.01295.x [DOI] [PubMed] [Google Scholar]
- Elek Z, Dauffy-Richard E, Gosselin F. (2010) Carabid species responses to hybrid poplar plantations in floodplains in France. Forest Ecology and Management 260:1446-1455. doi: 10.1016/j.foreco.2010.07.034 [DOI] [Google Scholar]
- Elek Z, Lövei GL. (2007) Patterns in ground beetle (Coleoptera: Carabidae) assemblages along an urbanisation gradient in Denmark. Acta Oecologica 32:104-111. doi: 10.1016/j.actao.2007.03.008 [DOI] [Google Scholar]
- Elron E, Shlagman A, Gasith A. (2007) First detailed report of predation on anuran metamorphs by terrestrial beetle larvae. Herpetological Reviews 38:30-33. [Google Scholar]
- Emmerich H, Thiele H-U. (1969) Wirkung von Farnesylmethyläther auf die Ovarienreifung von Pterostichus nigrita F. Naturwissenschaften 56: 641. doi: 10.1007/BF01185755 [DOI] [PubMed]
- Erbeling L, Paarmann W. (1986) The role of a circannual rhythm of thermoregulation on the control of the reproductive cycle of the desert carabid beetle Thermophilumsexmaculatum F. In: Den Boer PJ, Luff ML, Mossakowski D, Weber F (Eds) Carabid beetles - Their Adaptations and Dynamics. Gustav Fischer Verlag, Stuttgart-New York, 125–146.
- Ermakov AI. (2004) Structural changes in the carabid fauna of forest ecosystems under a toxic impact. Russian Journal of Ecology 35:403-408. doi: 10.1023/B:RUSE.0000046977.30889.a1 [DOI] [Google Scholar]
- Erwin TL. (1967) Bombardier beetles (Coleoptera, Carabidae) of North America: part II. Biology and behavior of Brachinus pallidus Erwin in California. Coleopterists Bulletin 21:41-55. [Google Scholar]
- Erwin TL. (1979a) A review of the Natural history and Evolution of Ectoparasitoid Relationships in Carabid Beetles. In: Erwin TL, Ball GE, Whitehead DR, Halpern AL (Eds) Carabid Beetles: Their Evolution, Natural history, and Classification. Dr. W. Junk Publishers, The Hague, 479–484.
- Erwin TL. (1979b) Thoughts on the evolutionary history of ground beetles: Hypothesis generated from comparative faunal analyses of lowland forest sites in temperate and tropical regions. In: Erwin TL, Ball GE, Whitehead DR (Eds) Carabid beetles, their evolution, natural history, and classification. W. Jung, The Hague, 539–592.
- Erwin TL. (1996) Arboreal beetles of neotropical forests: Agra Fabricius, the cayennensis complex (Coleoptera: Carabidae: Lebiini: Calleida). Annales Zoologici Fennici 33:17-21. [Google Scholar]
- Erwin TL. (2007) A treatise on the Western Hemisphere Caraboidea (Coleoptera). Their classification, distributions and ways of life. Vol. I (Trachypachidae, Carabidae – Nebriiformes 1). Pensoft Sofia-Moscow, 323 pp, 22 plates.
- Erwin TL, Ball GE, Whitehead DR, Halpern AL.Eds (1979) Carabid Beetles: Their Evolution, Natural history, and Classification. Dr. W. Junk Publishers, The Hague, 635 pp.
- Erwin TL, Geraci CJ. (2008) New genera of Western Hemisphere Pseudomorphini (Insecta, Coleoptera, Carabidae) with notes on their distributions, ways of life, and hypothesized relationships. In: Penev L, Erwin TL, Assmann T (Eds) Back to the roots and back to the future. Pensoft, Sofia, 77–100.
- Erwin TL, Pearson DL. (2008) A treatise on the Western Hemisphere Caraboidea (Coleoptera). Their classification, distributions and ways of life. Vol. II (Nebriiformes 2 - Cicindelidae). Pensoft, Sofia-Moscow, 365 pp, 33 plates.
- Evans MEG, Forsythe TG. (1985) Feeding mechanisms, and their variation in form, of some adult ground-betles (Coleoptera: Caraboidea). Acta Entomologica Sinica 206:113-143. [Google Scholar]
- Eversham BC, Roy DB, Telfer MG. (1996) Urban, industrial and other manmade sites as analogues of natural habitats for carabidae. Annales Zoologici Fennici 33:149-156. [Google Scholar]
- Eyre MD, Luff ML. (1990) A preliminary classification of European grassland habitats using carabid beetles. In: Stork NE (Ed) The role of groundbeetles in ecological and environmental studies. Intercept, Andover, Hampshire, 227–236.
- Fahrig L. (2003) Effects of habitat fragmentation on biodiversity. Annual Review of Ecology and Systematics 34:487-515. doi: 10.1146/annurev.ecolsys.34.011802.132419 [DOI] [Google Scholar]
- Falke B, Oevermann S, Assmann T. (2000) Ground beetles (Coleoptera, Carabidae) in a medieval wood-pasture reserve in north-west Germany. In: Brandmayr P, Lövei G, Zetto Brandmayr T, Casale A, Taglianti V (Eds) Natural history and applied ecology of carabid beetles, Pensoft, 265–275.
- Fawki S, Toft S. (2005) Food preferences and the value of animal food for the carabid beetle Amara similata (Gyll.) (Col., Carabidae). Journal of Applied Entomology 129:551-556. doi: 10.1111/j.1439-0418.2005.00992.x [DOI] [Google Scholar]
- Fedorenko DN. (1996) Reclassification of world Dyschirini, with a revision of the palearctic fauna (Coleoptera, Carabidae). Pensoft, Sofia-Moscow-St.Petersburg, 224 pp.
- Ferenz HJ. (1975) Anpassungen von Pterostichus nigrita F. (Coleoptera, Carabidae) an subartische Bedingungen. Oecologia 19:49-57. doi: 10.1007/BF00377589 [DOI] [PubMed] [Google Scholar]
- Ferenz HJ. (1977) Two-step photoperiodic and hormonal control of reproduction in the female beetle, Pterostichus nigrita. Journal of Insect Physiology 23:671-676. doi: 10.1016/0022-1910(77)90082-8 [DOI] [Google Scholar]
- Ferenz HJ, Hölters W. (1975) Corpus allatum hormone induced maturation in males of three carabid species of the genus Pterostichus (Col., Carab.). Entomologia Experimentalis et Applicata 18:238-243. doi: 10.1111/j.1570-7458.1975.tb02375.x [DOI] [Google Scholar]
- Ford ED. (2009) The importance of a research data statement and how to develop one. Annales Zoologici Fennici 46:82-92. [Google Scholar]
- Forel J, Leplat J. (1995) Les Carabes de France 1–2. Sciences Nat, Venette, 316 pp, maps, 57 plates.
- Forel J, Leplat J. (1998) Faune des Carabus de la peninsula ibérique. Magellanes 2:1-168. [Google Scholar]
- Forel J, Leplat J. (2001) Faune des carabiques de France -1. Magellanes 5:1-95. [Google Scholar]
- Forel J, Leplat J. (2003) Faune des carabiques de France -11 Magellanes 7: 1–159.
- Forel J, Leplat J. (2005) Faune des carabiques de France -10. Magellanes 12:1-129. [Google Scholar]
- Fournier E, Loreau M. (2001) Respective roles of recent hedges and forest patch remnants in the maintenance of ground-beetle (Coleoptera: Carabidae) diversity in an agricultural landscape. Landscape Ecology 16:17-32. doi: 10.1023/A:1008115516551 [DOI] [Google Scholar]
- Främbs H. (1990) Changes in carabid beetle populations on a regenerating, excavated peat bog in northwestern Germany. In: Stork NE (Ed) The role of ground beetles in ecological and environmental studies, Intercept, Andover, Hampshire, 157–169.
- Frank JH, Erwin TL, Hemenway RC. (2009) Economically Beneficial Ground Beetles. The specialized predators Pheropsophus aequinoctialis (L.) and Stenaptinus jessoensis (Morawitz): Their laboratory behavior and descriptions of immature stages (Coleoptera, Carabidae, Brachininae). Zookeys 14:1-36. doi: 10.3897/zookeys.14.188 [DOI] [Google Scholar]
- Freitag R, Kavanaugh DH. (1993) A new species of Cicindela (Cicindela) (Coleoptera: Carabidae: Cicindelini) from remnant native grassland in Santa Cruz County, California. Coleopterists Bulletin 47:113-120. [Google Scholar]
- Fuellhaas U. (2000) Restoration of degraded fen grassland - effects of long-term inundation and water logging on ground beetle populations (Coleoptera: Carabidae). In: Brandmayr P, Lövei G, Zetto Brandmayr T, Casale A, Taglianti V (Eds) Natural history and applied ecology of carabid beetles, Pensoft, 251–263.
- Fuller RJ, Oliver TH, Leather SR. (2008) Forest management effects on carabid beetle communities in coniferous and broadleaved forests: implications for conservation. Insect Conservation and Diversity 1:242-252. doi: 10.1111/j.1752-4598.2008.00032.x [DOI] [Google Scholar]
- GAC (1999) Laufkäfer in Auen. Angewandte Carabidologie – Supplement 1:1-144. [Google Scholar]
- GAC (2001) Laufkäfer im Wald. Angewandte Carabidologie – Supplement 2:1-122. [Google Scholar]
- GAC (2004) Laufkäfer in Xerothermbiotopen. Angewandte Carabidologie – Supplement 3:1-114. [Google Scholar]
- Ganglbauer L. (1892) Die Käfer von Mitteleuropa 1 - Familienreihe Caraboidea. Verlag Carl Gerold’s Sohn, Wien, 557 pp.
- Galián J, Prüser F, De la Rúa P, Serrano J, Mossakowski D. (1996) Cytological and molecular differences in the Ceroglossus chilensis species complex (Coleoptera: Carabidae). Annales Zoologici Fennici 33:23-30. [Google Scholar]
- Gärdenfors U (Ed) (2005) Rödlistade arter i Sverige 2005. The 2005 Red List of Swedish species. ArtDatabanken, Uppsala, 496 pp.
- Gardiner MM, Landis DA, Gratton C, Schmidt N, O'Neal M, Mueller E, Chacon J, Heimpel GE. (2010) Landscape composition influences the activity density of Carabidae and Arachnida in soybean fields. Biological Control 55:11-19. doi: 10.1016/j.biocontrol.2010.06.008 [DOI] [Google Scholar]
- Gaston KJ, Spicer JI. (2004) Biodiversity. An introduction. 2nd edition. Blackwell Publishing, Oxford, UK, 208 pp.
- Gaublomme E, Hendrickx F, Dhuyvetter H, Desender K. (2008) The effects of forest patch size and matrix type on changes in carabid beetle assemblages in an urbanized landscape. Biological Conservation 141:2585-2596. doi: 10.1016/j.biocon.2008.07.022 [DOI] [Google Scholar]
- Gebert J. (2006) Die Sandlaufkäfer und Laufkäfer von Sachsen. Beiträge zur Insektenfauna Sachsens, Band 4, Teil 1 (Carabidae: Cicindelini-Lorocerini): 1–180.
- Gilbert O. (1956) The natural history of four species of Calathus (Coleoptera, Carabidae) living on sand dunes in Anglesey, North Wales. Oikos 7:22-47. doi: 10.2307/3564982 [DOI] [Google Scholar]
- Grandchamp A-C, Bergamini A, Stofer S, Niemelä J, Duelli P, Scheidegger C. (2005) The influence of grassland management on ground beetles (Carabidae, Coleoptera) in Swiss montane meadows. Agriculture, Ecosystems and Environment 110:307-317. doi: 10.1016/j.agee.2005.04.018 [DOI] [Google Scholar]
- Greenslade PJM. (1964) Pitfall trapping as a method for studying populations of Carabidae (Coleoptera). Journal of Animal Ecology 33:301-310. doi: 10.2307/2632 [DOI] [Google Scholar]
- Griffiths GJK, Winder L, Holland JM, Thomas CFG, Williams E. (2007) The representation and functional composition of carabid and staphylinid beetles in different field boundary types at a farm-scale. Biological Conservation 135:145-152. doi: 10.1016/j.biocon.2006.09.016 [DOI] [Google Scholar]
- Gruttke H. (1994) Dispersal of carabid species along a linear sequence of young hedge plantations. In: Desender K, Dufrêne M, Loreau M, Luff ML, Maelfait J-P (Eds) Carabid beetles: ecology and evolution. Series Entomologica 51:229-303. [Google Scholar]
- Gueorguiev BV. (2007) Annotated catalogue of the carabid beetles of Albania (Coleoptera: Carabidae). Pensoft, Sofia-Moscow, 243 pp.
- Gueorguiev VB, Gueorguiev BV. (1995) Catalogue of the ground-beetles of Bulgaria (Coleoptera: Carabidae). Pensoft, Sofia-Moscow, 279 pp.
- Gueorguiev VB, Sakalian VP, Gueorguiev BV. (1997) Biogeography of the endemic ground-beetles (Coleoptera: Carabidae) in Bulgaria. Pensoft, Sofia-Moscow, 73 pp.
- Haberman H. (1968) Eesti jooksiklased. Zooloogia ja Botaanika Instituut, Tallinn, 598 pp.
- Habu A. (1967) Fauna Japonica. Carabidae. Truncatipennes Group. (Insecta: Coleoptera). Biogeographical Society of Japan. Tokyo Electrical Engineering College Press, pub. 338 pp, 27 plates.
- Habu A. (1973) Fauna Japonica. Carabidae: Harpalini (Insecta: Coleoptera). Keigaku Publishing Co Ltd. Tokyo, 430 pp, 21 plates.
- Habu A. (1978) Fauna Japonica. Carabidae: Platynini (Insecta: Coleoptera). Keigaku Publishing Co Ltd. Tokyo, 477 pp, 36 plates.
- Hammond PM. (1979) Wing-folding mechanisms of beetles, with special reference to investigations of adephagan phylogeny. In: Erwin TL, Ball GE, Whitehead DR, Halpern AL (Eds) Carabid Beetles: Their Evolution, Natural History, and Classification. Dr. W. Junk Publishers, The Hague, 113–180.
- Hance T. (1987) Predation impact of carabids at different population densities on Aphis fabae development in sugar beet. Pedobiologia 30:251-262. [Google Scholar]
- Hansen V. (1968) Biller XXIV – Sandspringere og lobebiller – Larverne ved Sb.G. Larsson. G-E-C Gads Vorlag, Kobenhavn, 449 pp.
- Haslett JR. (2007) European strategy for the conservation of invertebrates. Convention on the Conservation of European Wildlife and Natural Habitats (Bern Convention). Nature and environment, no. 145. Council of Europe Publishing, Strasbourg, 91 pp.
- Hatteland BA, Hauge E, Kirkendall LR, Solhøy T. (2005) Diversity and habitat preferences of ground beetles (Coleoptera, Carabidae) in a coastal area of North Trøndelag, Central Norway. DIAS Report 114:125-135. [Google Scholar]
- Heessen HJL. (1981) On the reproduction and survival in two ecologically related beetle-species of the forest floor. PhD-thesis Wageningen Agricultural University. Krips Repro Meppel, the Netherlands, 84 pp.
- Hemenway R, Whitcomb WH. (1967) Ground beetle of the Genus Lebia Latreille in Arkansas (Coleoptera: Carabidae): ecology and geographical distribution. Proceedings of Arkansas Academy of Science 21:15-20. [Google Scholar]
- Hendrickx F, Maelfait J-P, Desender K, Aviron S, Bailey D, Diekotter T, Lens L, Liira J, Schweiger O, Speelmans M, Vandomme V, Butger R. (2009). Pervasive effects of dispersal imitation on within- and among-community species richness in agricultural landscapes. Global Ecology and Biogeography 18:607-616. doi: 10.1111/j.1466-8238.2009.00473.x [DOI] [Google Scholar]
- Hendrickx F, Maelfait JP, Van Wingerden W, Schweiger O, Speelmans M, Aviron S, Augenstein I, Billeter R, Bailey D, Bukacek R, Burel F, Diekotter T, Dirksen J, Herzog F, Liira J, Roubalova M, Vandomme V, Bugter R. (2007) How landscape structure, land-use intensity and habitat diversity affect components of total arthropod diversity in agricultural landscapes. Journal of Applied Ecology 44:340-351. doi: 10.1111/j.1365-2664.2006.01270.x [DOI] [Google Scholar]
- Hengeveld R. (1980a) Food specialization in ground beetles: an ecological or a phylogenetical process? (Coleoptera, Carabidae). Netherlands Journal of Zoology 30:585-594. doi: 10.1163/002829679X00205 [DOI] [Google Scholar]
- Hengeveld R. (1980b) Qualitative and quantitative aspects of the food of ground beetles (Coleoptera, Carabidae). Netherlands Journal of Zoology 30:557-563. [Google Scholar]
- Hengeveld R. (1987) Scales of variation: their distinction and ecological importance. Annales Zoologici Fennici 24:195-202. [Google Scholar]
- Herrera L, Arricibita FC. (1990) Los carabidos de Navarra España (Coleoptera, Carabidae). Brill, Leiden, Entomonograph 12:1-241. [Google Scholar]
- Heydemann B. (1962) Die biozönotische Entwicklung vom Vorland zum Koog. II. Teil: Käfer (Coleoptera). Akademie der Wissenschaften und der Literatur, Mainz - Abhandlungen der Mathematisch-Naturwissenschaftlichen Klasse 11:765-964. [Google Scholar]
- Hieke F, Wrase DW. (1988) Faunistik der Laufkäfer Bulgariens (Coleoptera, Carabidae): Deutsche Entomologische Zeitschrift 35: 1–171. doi: 10.1002/mmnd.19880350102 [DOI]
- Hoffmann HJ. (1969) Neuro-endocrine control of diapause oocyte maturation in the beetle Pterostichus nigrita. Journal of Insect Physiology 16:629-642. doi: 10.1016/0022-1910(70)90096-X [DOI] [Google Scholar]
- Holdhaus K, Lindroth CH. (1939) Die europäischen Carabiden mit boreoalpiner Verbreitung. Annalen des Nasturhistorischen Museums, Wien 50:123-293. [Google Scholar]
- Holland JM. (Ed) (2002) The Agroecology of Carabid Beetles. Intercept Ltd, Hampshire, UK, 356 pp.
- Hölldobler B, Wilson EO. (1990) The Ants. Springer Verlag, Berlin.
- Hollmen A, Välimäki P, Itämies J, Oksanen J. (2008) The value of open power line habitat in conservation of ground beetles (Coleoptera: Carabidae) associated with mires. Journal of Insect Conservation 12:163-177. doi: 10.1007/s10841-007-9076-7 [DOI] [Google Scholar]
- Honek A, Martinkova Z, Jarosik V. (2003) Ground beetles (Carabidae) as seed predators. European Journal of Entomology 100:531-544. [Google Scholar]
- Honek A, Martinkova Z, Saska P. (2005) Post-dispersal predation of Taraxacum officinale (dandelion) seed. Journal of Ecology 93:345-352. doi: 10.1111/j.1365-2745.2005.00987.x [DOI] [Google Scholar]
- Honek A, Martinkova Z, Saska P. (2011) Effect of size, taxonomic affiliation and geographic origin of dandelion (Taraxacum agg.) seeds on predation by ground beetles (Carabidae, Coleoptera). Basic and Applied Ecology 12:89-96. doi: 10.1016/j.baae.2010.11.003 [DOI] [Google Scholar]
- Honek A, Martinkova Z, Saska P, Pekar S. (2007) Size and taxonomic constraints determine the seed preferences of Carabidae (Coleoptera). Basic and Applied Ecology 8:343-353. doi: 10.1016/j.baae.2006.07.002 [DOI] [Google Scholar]
- Honek A, Saska P, Martinkova Z. (2006) Seasonal variation in seed predation by adult carabid beetles. Entomologia Experimentalis et Applicata 118:157-162. doi: 10.1111/j.1570-7458.2006.00376.x [DOI] [Google Scholar]
- Hutchinson EG. (1965) The ecologial theatre and the evolutionary play. Yale University Press, New Haven, USA.
- Huusela-Veistola E. (1996) Effects of pesticide use and cultivation techniques on ground beetles (Col., Carabidae) in cereal fields. Annales Zoologici Fennici 33:197-205. [Google Scholar]
- Hurka K. (1996) Carabidae of the Czech and Slovak Republics – Illustrated key: Kabourek, Zlín, 1–565.
- Hurka K, Jarošík V. (2003) Larval omnivory in Amara aenea (Coleoptera: Carabidae). European Journal of Entomology 100:329-335. [Google Scholar]
- Hurlbert SH. (1984) Pseudoreplication and the design of ecological field experiments. Ecological Monographs 54:187-211. doi: 10.2307/1942661 [DOI] [Google Scholar]
- Hyman PS, Parsons MS. (1992) A review of the scarce and threatened Coleoptera in Great Britain, Part 1 - Carabidae. The UK Joint Nature Conservation Committee, Peterborough: 99–155.
- Iablokov-Khnzorian SM. (1976) Fauna of the Armenian SSR. Coleopterous insects, the Ground-beetles (Carabidae). Izd-vo Akademia Nauk Armenia, Yerevan, 297 pp.
- Imura Y. (2002) Classification of the subtribe Carabina (Coleoptera, Carabidae) based on molecular phylogeny. Elytra (Tokyo) 30:1-28. [Google Scholar]
- Ishikawa R. (1984) Phylogeny and subgeneric classification of the genus Chaetocarabus (Coleoptera, Carabidae). Kontyu (Tokio) 53:94-109. [Google Scholar]
- Ishitani M, Kotze DJ, Niemelä J. (2003) Changes in carabid beetle assemblages across and urban-rural gradient in Japan. Ecography 26:481-489. doi: 10.1034/j.1600-0587.2003.03436.x [DOI] [Google Scholar]
- Janssen P, Fortin D and Hebert C. (2009) Beetle diversity in a matrix of old-growth boreal forest: influence of habitat heterogeneity at multiple scales. Ecography 32:423-432. doi: 10.1111/j.1600-0587.2008.05671.x [DOI] [Google Scholar]
- Jeannel R. (1926–1928) Monographie des Trechinae 1–3. Abeille, Paris, 32, 33, 35 : 221–550, 1–592, 1–808.
- Jeannel R. (1941–1942, 1949) Faune de France 39–40, 51: Coleoptere Carabiques 1, 2 supplement 1. Lechevalier et Fils, Paris, 1173 pp.
- Jonsson BG, Jonsell M. (1999) Exploring potential biodiversity indicators in boreal forests. Biodiversity and Conservation 8:1417-1433. doi: 10.1023/A:1008900309571 [DOI] [Google Scholar]
- Jørgensen HB, Toft S. (1997a) Food preference, diet dependent fecundity and larval development in Harpalus rufipes (Coleoptera: Carabidae). Pedobiologia 41:307-315. [Google Scholar]
- Jørgensen HB, Toft S. (1997b) Role of granivory and insectivory in the life cycle of the carabid beetle Amara similata. Ecological Entomology 22:7-15. doi: 10.1046/j.1365-2311.1997.00045.x [DOI] [Google Scholar]
- Juberthie C. (1969) Relations entre le climat, le microclimat et les Aphenops cerberus dans la Grotte de Sainte-Catherine. Annales de Spéléologie 24:75-104. [Google Scholar]
- Juberthie C. (1979) L’evolution des coléoptères Trechinae souterrains (Coleoptera, Carabidae. In: Den Boer PJ, Thiele HU, Weber F (Eds) On the Evolution of Behaviour in Carabid Beetles. Miscellaneous Papers, 18, Landbouwhogeschool Wageningen, 83–99.
- Juliano SA. (1985) Habitat associations, resources, and predators of Brachinus (Coleoptera: Carabidae) from southeastern Arizona. Canadian Journal of Zoology 63:1683-1691. doi: 10.1139/z85-250 [DOI] [Google Scholar]
- Kålås JA, Viken Å, Bakken T.Eds (2006) Norsk Rødliste 2006. The Norwegian Red List 2006. Artsdatabanken, Trondheim, Norway, 416 pp.
- Kamer N, Brückner M, Düring A, Mossakowski D. (2002) ND5 gene tree and the phylogenetic history of supertribe Carabitae. In: Szyszko J, Den Boer PJ, Bauer T (Eds) How to protect or what we know about carabid beetles. Warsaw Agricultural University Press, Warsaw, 331–335.
- Kamer N, Dormann W, Mossakowski D. (2008) Patterns of molecular variability in carabid beetles mostly from the Baltric Sea coast. In: Penev L, Erwin TL, Assmann T (Eds) Back to the roots and back to the future. Pensoft, Sofia, 195–206.
- Kane TC, Norton RM, Poulson TL. (1975) The ecology of a predaceous troglobitic beetle, Neoaphaenops tellkampfi (Coleoptera, Trechinidae). II Adult seasonality, feeding and recruitment. International Journal of Speleology 7:55-64. [Google Scholar]
- Kaufmann T. (1971) Hibernation in the Arctic Arctic beetle Pterostichus brevicornis, in Alaska. Journal of the Kansas Entomological Society 44:81-92. [Google Scholar]
- Kavanaugh DH. (1996) Phylogenetic relationships of the genus Pelophila Dejean to other basal grade Carabidae (Coleoptera). Annales Zoologici Fennici 33:31-37. [Google Scholar]
- Kavanaugh DH. (1998) Investigations of phylogenetic relationships among some basal grade Carabidae (Coleoptera): a report on work in progress. In: Ball GE, Casale A, Vigna Taglianti A (Eds) Phylogeny and classification of Caraboidea (Coleoptera: Adephaga). Museo Regionale di Scienze Naturali, Torino, 329–341.
- Keller I, Nentwig W, Largiadèr R. (2004) Recent habitat fragmentation due to roads can lead to significant genetic differentiation in an abundant flightless ground beetle. Molecular Ecology 13:2983-2994. doi: 10.1111/j.1365-294X.2004.02310.x [DOI] [PubMed] [Google Scholar]
- Kinnunen H, Järveläinen K, Pakkala T, Tiainen J. (1996) The effect of isolation on the occurrence of farmland carabids in a fragmented landscape. Annales Zoologici Fennici 33:165-171. [Google Scholar]
- Kinnunen H, Tiainen J, Tukia H. (2001) Farmland carabid beetle communities at multiple levels of spatial scale. Ecography 24:189-197. doi: 10.1034/j.1600-0587.2001.240209.x [DOI] [Google Scholar]
- Kirby P. (1992) Habitat management for invertebrates: a practical handbook. Sandy, Bedfordshire.
- Kjellsson G. (1985) Seed fate in a population of Carex pilulifera L. 2. Seed predation and its consequences for dispersal and seed bank. Oecologia 67:424-429. doi: 10.1007/BF00384950 [DOI] [PubMed] [Google Scholar]
- Klausnitzer B, Richter K. (1983) Presence of an urban gradient demonstrated for carabid associations. Oecologia 59:79-82. doi: 10.1007/BF00388077 [DOI] [PubMed] [Google Scholar]
- Klimeš P, Saska P. (2010) Larval and adult seed consumption affected by the degree of food specialization in Amara (Coleoptera: Carabidae). Journal of Applied Entomology 134:659-666. [Google Scholar]
- Koch D. (1986) Morphological-physiological studies on Pterostichus “nigrita” (Col., Carabidae), a complex of sibling species. In: Den Boer P, Luff ML, Mossakowski D, Weber F (Eds) Carabid Beetles, their adaptations and dynamics. Gustav Fischer, Stuttgart, 267–279.
- Kocher L. (1963) Catalogue commenté des Coléoptères du Maroc. I. Carabiques. Travaux de l’’Institut Scientifique Chérifien, Zoologie 27, Rabat, 170 pp.
- Koivula M. (2002a) The forest road network - a landscape element affecting the distribution of boreal carabid beetles (Coleoptera, Carabidae). In: Szyszko J, Den Boer PJ, Bauer T (Eds) How to protect or what we know about carabid beetles, Warsaw Agricultural University Press, Warsaw, 287–299.
- Koivula M. (2002b) Alternative harvesting methods and boreal carabid beetles (Coleoptera, Carabidae). Forest Ecology and Management 167:103-121. doi: 10.1016/S0378-1127(01)00717-4 [DOI] [Google Scholar]
- Koivula MJ. (2005) Effects of forest roads on spatial distribution of boreal carabid beetles (Coleoptera: Carabidae). The Coleopterists Bulletin 59:465-487. doi: 10.1649/815.1 [DOI] [Google Scholar]
- Koivula MJ. (2011) Useful model organisms, indicators, or both? Ground beetles (Coleoptera, Carabidae) reflecting environmental conditions. In: Kotze DJ, Assmann T, Noordijk J, Turin H, Vermeulen R (Eds) Carabid Beetles as Bioindicators: Biogeographical, Ecological and Environmental Studies. ZooKeys 100:287-317. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Koivula MJ, Kotze DJ. (2005) Carabid beetles in median strips of three highways around the city of Helsinki, Finland. DIAS Report 114:151-161. [Google Scholar]
- Koivula MJ, Kotze DJ, Salokannel J. (2005) Beetles (Coleoptera) in central reservations of three highway roads around the city of Helsinki, Finland. Annales Zoologici Fennici 42:615-626. [Google Scholar]
- Koivula M, Kukkonen J, Niemelä J. (2002) Boreal carabid-beetle (Coleoptera, Carabidae) assemblages along the clear-cut originated succession gradient. Biodiversity and Conservation 11:1269-1288. doi: 10.1023/A:1016018702894 [DOI] [Google Scholar]
- Koivula M, Spence JR. (2006) Effects of post-fire salvage logging on boreal mixed-wood ground beetle assemblages (Coleoptera, Carabidae). Forest Ecology and Management 236:102-112. doi: 10.1016/j.foreco.2006.09.004 [DOI] [Google Scholar]
- Koivula MJ, Vermeulen HJW. (2005) Highways and forest fragmentation – effects on carabid beetles (Coleoptera, Carabidae). Landscape Ecology 20:911-926. doi: 10.1007/s10980-005-7301-x [DOI] [Google Scholar]
- Kotze DJ. (2000) Ground beetle diversity patterns across Afro-montane forest/grassland ecotones in KwaZulu-Natal, South-. Africa. In: Brandmayr P, Lövei G, Zetto Brandmayr T, Casale A, Taglianti V (Eds) Natural history and applied ecology of carabid beetles, Pensoft, 231–240.
- Kotze DJ. (2008) The occurrence and distribution of carabid beetles (Carabidae) on islands in the Baltic Sea: a review. Journal of Insect Conservation 12:265-276. doi: 10.1007/s10841-008-9147-4 [DOI] [Google Scholar]
- Kotze DJ, Niemelä J. (2002) Year-to-year variation in carabid beetle (Coleoptera, Carabidae) assemblages on the Åland Islands, south-west Finland. Journal of Biogeography 29:375-386. doi: 10.1046/j.1365-2699.2002.00681.x [DOI] [Google Scholar]
- Kotze DJ, Niemelä J, Nieminen M. (2000) Colonization success of carabid beetles on Baltic islands. Journal of Biogeography 27:807-819. doi: 10.1046/j.1365-2699.2000.00456.x [DOI] [Google Scholar]
- Kotze DJ, O’Hara RB. (2003) Species decline - but why? Explanations of carabid beetle (Coleoptera, Carabidae) declines in Europe. Oecologia 135:138-148. [DOI] [PubMed] [Google Scholar]
- Krehan I. (1970) Die Steuerung der von Jahresrhythmik und Diapause bei Larval- und Imagoüberwinterern der Gattung Pterostichus (Coleoptera, Carabidae). Oecologia 6:58-105. doi: 10.1007/BF00345223 [DOI] [PubMed] [Google Scholar]
- Kromp B. (1999) Carabid beetles in sustainable agriculture: a review on pest control efficacy, cultivation impacts and enhancement. Agriculture, Ecosystems and Environment 74:187-228. doi: 10.1016/S0167-8809(99)00037-7 [DOI] [Google Scholar]
- Kryzhanovskij OL. (1983) Zhuki podotryada Adephaga: semeistva Rhysodidae, Trachypachidae), semeistvo Carabidae (vvodna chast’I obzor fauny SSSR). (The ground-beetles of the sunorder Adephaga: Families Rhysodidae, Trachypachidae, family Carabidae) – Introduction and review of the Russian fauna. Fauna SSSR, Leningrad, Nauka 1: 1–3443 (In Russian).
- Kryzhanovskij OL, Belousov IA, Kabak II, Kataev BM, Makarov KV, Shilenkov VG. (1995) A checklist of the groundbeetles of Russia and adjacent lands (Insecta, Coleoptera, Carabidae). Pensoft, Sofia-Moscow, 271 pp.
- Kult K. (1947) Key to the beetle family Carabidae of the Czechoslovak Repuplic. Entomologicke prirucky – Czechoslovak Entomological Society, Prague, 199 pp.
- Läärä E. (2009) Statistics: reasoning on uncertainty, and the insignificance of testing null. Annales Zoologici Fennici 46:138-157. [Google Scholar]
- Landres PB, Verner J, Thomas JW. (1988) Ecological uses of vertebrate indicator species: a critique. Conservation Biology 2:316-328. doi: 10.1111/j.1523-1739.1988.tb00195.x [DOI] [Google Scholar]
- Lange M, Gossner, MM, Weisser WW. (2011) Effect of pitfall trap type and diameter on vertebrate by-catches and ground beetle (Coleoptera: Carabidae) and spider (Araneae) sampling. Methods in Ecology and Evolution, in press. doi: 10.1111/j.2041-210X.2010.00062.x [DOI]
- Langor DW, Spence JR. (2006) Arthropods as ecological indicators of sustainability in Canadian forests. Forestry Chronicle 82:344-350. [Google Scholar]
- Larochelle A. (1990) The food of carabid beetles (Coleoptera: Carabidae, including Cicindelinae). Fabreries Supplément 5:1-132. [Google Scholar]
- Larochelle A, Larivière M-C. (2003) A natural history of the ground-beetles (Coleoptera: Carabidae) of America north of Mexico. Pensoft, Sofia-Moscow, 583 pp.
- Larsson SG. (1939) Entwicklungstypen und Entwicklungszeiten der dänischen Carabiden. Kobenhavn: 1–360, tab.
- Larsson SG, Gigja G. (1959) Coleoptera 1 - Synopsis of the species. Zoology of Iceland, 3 (46a): 1–218.
- Lawrence JF, Moore BP, Pyke JE, Weir TA. (1987). Zoological Catalogue of Australia - Coleoptera: Archostemata, Myxophaga, Adephaga. Australian Governement Publishing Service: viii + 44: 4 pp. [Google Scholar]
- Leather SR, Basset Y, Hawkins BA. (2008) Insect conservation: finding the way forward. Insect Conservation and Diversity 1:67-69. doi: 10.1111/j.1752-4598.2007.00005.x [DOI] [Google Scholar]
- Ledoux G, Roux Ph. (2005) Nebria (Coleoptera, Nebriidae) Faune mondiale. Société Linéenne de Lyon, Lyon, 976 pp.
- Levins R. (1970) Extinction. Lectures Notes in Mathematics 2:75-107. [Google Scholar]
- Lewis OT, New TR, Stewart AJA. (2007) Insect conservation: progress and prospects. In: Stewart AJA, New TR, Lewis OT (Eds) Insect conservation biology. The Royal Entomological Society / CABI, Wallingford, 431–436. doi: 10.1079/9781845932541.0431 [DOI]
- Liebherr JK. (1986) The Cochise filter barrier: the major vicariant mechanism in the Agonum extensicolle species group (Carabidae: Platynini). In: Den Boer P, Luff ML, Mossakowski D, Weber F (Eds) Carabid Beetles, their adaptations and dynamics. Gustav Fischer, Stuttgart, 255–266.
- Liebherr JK, Will KW. (1998) Inferring phylogenetic relationships within the Carabidae (Insecta, Coleoptera) from characters of the female reproductive tract. In: Ball GE, Casale A, Vigna Taglianti A (Eds) Phylogeny and classification of Caraboidea (Coleoptera: Adephaga). Museo Regionale di Scienze Naturali, Torino, 107–170.
- Lindenmayer DB, Margules CR, Botkin DB. (2000) Indicators of biodiversity for ecologically sustainable forest management. Conservation Biology 14:941-950. doi: 10.1046/j.1523-1739.2000.98533.x [DOI] [Google Scholar]
- Lindroth CH. (1931) Die Insektenfauna Islands und ihre Probleme. Zoologiske Bidrag, Uppsala 13:105-218. [Google Scholar]
- Lindroth CH. (1945a, b, 1949) Die Fennoscandischen Carabidae: I Spezieller Teil, II Die Karten, III Algemeiner Teil. Göteborg: 1–709, 1–277, 1–911.
- Lindroth CH. (1956) A revision of the genus Synuchus. Transactions of the Royal Entomological Society, London 108:485-576. doi: 10.1111/j.1365-2311.1956.tb01274.x [DOI] [Google Scholar]
- Lindroth CH. (1960) Catalogus Coleopterorum Fennoscandiae et Daniae. Auctoribus V. Hanssen, E. Klefbeck, O Sjöberg, G. Stenius & A. Strand, redigenda curavit Carl H. Lindroth. Entomologiska Sällskaper, Lund, 44 pp.
- Lindroth CH. (1961–1969) The ground-beetles of Canada and Alaska. Opuscula Entomologica Suppl. 20, 24, 29, 33–35. Entomologiska Sällskapet, Lund.
- Lindroth CH. (1985–1986) The Carabidae (Coleoptera) of Fennoscandia and Denmark. Fauna Entomologica Scandinavica. Volume 15, Part I, II. Scandinavian Science Press, Ltd. Leiden.
- Lindroth CH. (1988, 1992a, b) Ground Beetles (Carabidae) of Fennoscandia – A Zoogeographical Study. Translation by J. Adis, Erwin TL of Lindroth CH (1945, 1949). Die Fennoskandischen Carabidae. Smithsonian Institution: I: 1–630 (1992), II: 1–271 (1988), III: 1–814 (1992).
- Löbl I, Smetana A.Eds (2003) Catalogue of Palaearctic Coleoptera, Vol. 1. Archostemata – Myxophaga – Adephaga: Apollo Books, Stenstrup, 819 pp.
- Loreau M. (1990) The significance of Intra- and Interspecific Competition in Carabid beetles. In Stork N (Ed), The Role of Ground Beetles in Ecological and Environmental studies. Intercept, Adover, Hampshire, 31–37.
- Lorenz W. (1998) Systematic list of the extant ground beetles of the world (Insecta. Coleoptera “Geadephaga: Trachypachidae and Carabidae incl. Paussinae, Cicindelinae, Rhysodinae). Lorenz, Tutzing, 502 pp.
- Lorenz W. (2005) Nomina Carabidarum. A directory of the scientific names of ground beetles (Insecta. Coleoptera “Geadephaga: Trachypachidae and Carabidae incl. Paussinae, Cicindelinae, Rhysodinae). Lorenz, Tutzing, 993 pp.
- Löser S. (1970) Brutfürsorge und Brutpflege bei Laufkäfern der Gattung Abax. Verhandlungen Der Deutschen Zoologischen Gesellschaft, Würzburg 1969:322-326. [Google Scholar]
- Lövei GL, Magura T. (2006) Body size changes in ground beetle assemblages - a re-analysis of Braun et al. (2004)’s data. Ecological Entomology 31:411-414. [Google Scholar]
- Lövei GL, Magura T. (2011) Can carabidologists spot a pitfall? The non-equivalence of two components of sampling effort in pitfall-trapped ground beetles (Carabidae). Community Ecology, in press.
- Lövei GL, Sunderland KD. (1996) Ecology and behavior of ground beetles (Coleoptera: Carabidae). Annual Review of Entomology 41:231-256. doi: 10.1146/annurev.en.41.010196.001311 [DOI] [PubMed] [Google Scholar]
- Lövei GL, Toft S.Eds (2005) European Carabidology – Proceedings of the 11th European Carabidologists’ Meeting. Danish Institute of Agricultural Sciences 114: 40: 1 pp. [Google Scholar]
- Luff ML. (1993) The Carabidae (Coleoptera) larvae of Fennoscandia and Denmark. Fauna Entomologica Scandinavica 28:1-186. [Google Scholar]
- Luff ML. (1998) Provisional atlas of the ground beetles (Coleoptera Carabidae) of Britain. Institute for Terrestrial Ecology, Monks Wood, 194 pp.
- Luff ML. (2002) Carabid assemblage organization and species composition. In: Holland JM (Ed) The Agroecology of Carabid Beetles, Intercept Limited, Hampshire, UK, 41–79.
- Luff ML. (2007) The Carabidae (ground beetles) of Britain and Ireland. RES Handbook Volume 4, Part 2. (2nd edition), Field Studies Council, Shrewsbury, 247 pp.
- Luff ML, Eyre MD, Rushton SP. (1989) Classification and ordination of habitats of ground beetles (Coleoptera, Carabidae) in north-east England. Journal of Biogeography 16:121-130. doi: 10.2307/2845086 [DOI] [Google Scholar]
- Luigioni P. (1929) I Coleotteri d’Italia. Memorie Accad. pont Nuovi Lincei, Roma, 1160 pp.
- Luka H, Marggi W, Huber C, Gonseth Y, Nagel P. (2009) Fauna Helvetica 24 – Carabidae. Centre Suisse de Cartographie de la Faune, 677 pp.
- Lund RD, Turpin FT. (1977) Carabid damage to weed seeds found in Indiana cornfields. Environmental Entomology 6:695-698. [Google Scholar]
- Lundgren JG, Lehman RM. (2010) Bacterial Gut Symbionts Contribute to Seed Digestion in an Omnivorous Beetle. PLoS ONE 5: e10831. doi: 10.1371/journal.pone.0010831 [DOI] [PMC free article] [PubMed]
- Lundgren JG, Rosentrater KA. (2007) The strength of seeds and their destruction by granivorous insects. Arthropod-Plant Interactions 1:93-99. doi: 10.1007/s11829-007-9008-1 [DOI] [Google Scholar]
- MacArthur RH, Wilson EO. (1967) The theory of island biogeography. Monographs in Population Biology, Princeton University Press, New York, Princeton, NJ, 203 pp.
- Machado A. (1992) Monografia de los carabidos de las islas Canarias. Instituto de Estudios Canarios, La Laguna, 734 pp.
- Maddison DR, Baker MD, Ober KA. (1998) A preliminary phylogenetic analysis of 18S ribosomal DNA of carabid beetles (Coleoptera). In: Ball GE, Casale A, Vigna Taglianti A (Eds) Phylogeny and classification of Caraboidea (Coleoptera: Adephaga). Museo Regionale di Scienze Naturali, Torino, 229–250.
- Maddison DR, Moore W, Baker MD, Ellis TM, Ober KA, Cannone JJ, Gutell RR. (2009) Monophyly of terrestrial adephagan beetles as indicated by three nuclear genes (Coleoptera: Carabidae and Trachypachidae). Zoologica Scripta 38:43-62. doi: 10.1111/j.1463-6409.2008.00359.x [DOI] [PMC free article] [PubMed] [Google Scholar]
- Mader H-J. (1984) Animal habitat isolation by roads and agricultural fields. Biological Conservation 29:81-96. doi: 10.1016/0006-3207(84)90015-6 [DOI] [Google Scholar]
- Maelfait J-P, Desender K, Dufrêne M. (1994) Carabid beetles and nature conservation research in Belgium: a review. In: Desender K, Dufrêne M, Loreau M, Luff ML, Maelfait J-P (Eds) Carabid beetles: ecology and evolution. Series Entomologica 51:319-323. [Google Scholar]
- Magistretti M. (1965) Fauna d’Italia, Cicindelidae, Carabidae – Catalogo topografico. Edizione Calderini, Bologna, 512 pp.
- Magura T, Koedoeboecz V, Tóthmérész B. (2001) Effects of habitat fragmentation on carabids in forest patches. Journal of Biogeography 28: 129–138. doi: 10.1046/j.1365-2699.2001.00534.x [DOI]
- Magura T, Lövei GL, Tóthmérész B. (2010) Does urbanization decrease diversity in ground beetle (Carabidae) assemblages? Global Ecology and Biogeography 19: 16–26. doi: 10.1111/j.1466-8238.2009.00499.x [DOI]
- Magura T, Tóthmérész B, Elek Z. (2006) Changes in carabid beetle assemblages as Norway spruce plantation age. Community Ecology 7:1-12. doi: 10.1556/ComEc.7.2006.1.1 [DOI] [Google Scholar]
- Mahler V, Suikat R, Assmann T. (1996) VII. Red List of beetles of the Wadden Sea area. Helgoländer Meeresuntersuchungen 50:83-96. doi: 10.1007/BF02366176 [DOI] [Google Scholar]
- Makarov KV, Bokhovko EE. (2005) Continuity of chaetom pattern in Brachinus-larvae developing with hypermetamorphosis (Coleoptera: Carabidae). Russian Entomological Journal 14:263-274. [Google Scholar]
- Mandl K. (1972) Coleoptera: Cicindelidae und Carabidae-Carabinae. Catalogus Faunae Austriae 15a, Wien, 1–16.
- Mandl K. (1978) Coleoptera: Carabinae II. Catalogus Faunae Austriae 15b, Wien, 1–58.
- Marggi WA. (1992) Faunistik der Sandleufkäfer und Laukäfer der Schweiz (Cicindelidae & Carabidae - Teil 1 / Text, Teil 2 / Verbreitungskarten. CSCF: 1–477, 1–243.
- Martínez-Navarro EM, Galián J, Serrano J. (2005) Phylogenetic relationships among subtribes of Harpalini Bonelli (Coleoptera, Carabidae) inferred from mitochondrial DNA sequences. In: Lövei G, Toft S (Eds) European Carabidology 2003 (Proceedings of the 11th European Carabidologists' Meeting). DIAS Report Plant Production, No. 114, Danish Institute of Agricultural Sciences, 219–230.
- Martínez-Navarro EM, Serrano J, Galián J. (2011) Chromosomes of Trachypachus Motschulsky and the evolution of the ancestral adephagan karyotype (Coleoptera). Journal of Zoological Systematics and Evolutionary Research, doi: 10.1111/j.1439–0469.2010.00613.x, in press.
- Martins da Silva P, Aguiar CAS, Niemelä J, Sousa JP, Serrano ARM. (2008) Diversity patterns of ground-beetles (Coleoptera: Carabidae) along a gradient of land-use disturbance. Agriculture, Ecosystems and Environment 124:270-274. doi: 10.1016/j.agee.2007.10.007 [DOI] [Google Scholar]
- Maryański M, Kramarz P, Laskowski R, Niklińska M. (2002) Decreased energetic reserves, morphological changes and accumulation of metals in carabid beetles (Poecilus cupreus L.) exposed to Zinc- and Cadmium-contaminated food. Ecotoxicology 11:127-139. doi: 10.1023/A:1014425113481 [DOI] [PubMed] [Google Scholar]
- Matalin A. (1994) The strategy of dispersal behaviour in some Carabidae species of Southeastern Europe. In: Desender K, Dufrêne M, Loreau M, Luff ML, Maelfait J-P (Eds) Carabid beetles: Ecology and Evolution.Kluwer Academic Publishers, Dordrecht, The Netherlands, 183–188.
- Matalin AV. (1997a) Specific features of life cycle of Pseudoophonus (s. str.) rufipes Deg. (Coleoptera, Carabidae) in Southwest Moldova. The Biological Bulletin 24:371-381. [Google Scholar]
- Matalin AV. (1997b) Life cycles of carabids of the genus Stenolophus (Coleoptera, Carabidae) in the steppe zone of Europe. Entomological Review 77:1181-1190. [Google Scholar]
- Matalin AV. (1997c) Peculiarities of spatial and temporal differentiation of carabids (Coleoptera, Carabidae) in the steppe zone. Entomological Review 77:1155-1166. [Google Scholar]
- Matalin AV. (1998) Polyvariance of Harpalus (s. str.) affinis Schrank (Coleoptera, Müller, H.J. (1970). Formen der Dormanz bei Insekten. Nova Acta Leopoldina 35:7-25. [Google Scholar]
- Matalin A. (2003) Variations in flight ability with sex and age in ground beetles (Coleoptera, Carabidae) of south-western Moldova. Pedobiologia47: 311–319. doi: 10.1078/0031-4056-00195 [DOI]
- Matalin AV. (2008) Evolution of biennal life cycles in ground beetles (Coleoptera, Carabidae) of the Western Palaeartic. In: Penev L, Erwin T, Assmann T (Eds) Back to the roots and back to the future. Proceedings of XIII European Carabidologist Meeting, Blagoevgrad, 259–284.
- Matveinen-Huju K, Niemelä J, Rita H, O’Hara RB. (2006) Retention-tree groups in clear-cuts: Do they constitute ”life-boats” for spiders and carabids? Forest Ecology and Management 230: 119–135. doi: 10.1016/j.foreco.2006.04.025 [DOI]
- Mccracken DI. (1994) A fuzzy classification of moorland ground beetles and (Coleoptera, Carabidae) and plant communities. Pedobiologia 18:12-27. [Google Scholar]
- McFerran DM, Meharg MJ, Montgomery WI, McAdam JH. (1994) The impact of grazing on communities of ground-dwelling beetles (Coleoptera, Carabidae) in upland vegetation in north-east Ireland. In: Desender K, Dufrêne M, Loreau M, Luff ML, Maelfait J-P (Eds) Carabid beetles: ecology and evolution. Series Entomologica 51:325-330. [Google Scholar]
- Melis C, Olsen CB, Hyllvang M, Gobbi M, Stokke BG, Røskaft E. (2010) The effect of traffic intensity on ground beetle (Coleoptera: Carabidae) assemblages in central Sweden. Journal of Insect Conservation 14:159-168. doi: 10.1007/s10841-009-9240-3 [DOI] [Google Scholar]
- Merivee E, Ploomi A, Milius M, Luik A, Heidemaa M. (2005) Electrophysiological identification of antennal pH receptors in the ground beetle Pterostichus oblongopunctatus. Physiological Entomology 30:122-133. doi: 10.1111/j.1365-3032.2005.00435.x [DOI] [Google Scholar]
- Milius M, Merivee E, Williams I, Luik A, Mänd M, Must A. (2006) A new method for electrophysiological identification of antennal pH receptor cells in ground beetles: The example of Pterostichus aethiops (Panzer, 1796) (Copeoptera, Carabidae. Journal of Insect Physiology 52:960-967. doi: 10.1016/j.jinsphys.2006.06.003 [DOI] [PubMed] [Google Scholar]
- Millán de la Peña N, Butet A, Delettre Y, Morant P, Burel F. (2003) Landscape context and carabid beetles (Coleoptera: Carabidae) communities of hedgerows in western France. Agriculture, Ecosystems and Environment 94:59-72. doi: 10.1016/S0167-8809(02)00012-9 [DOI] [Google Scholar]
- Mlynar Z. (1977) Revision der Arten und Unterarten der Gattung Molops Bonelli (Coleoptera, Carabidae. Folia entomologica Hungarica 30:3-150. [Google Scholar]
- Moore BP. (1971) Notes on an extraordinary blood-feeding carabid larva (Coleoptera) from Port Elizabeth. Annales of Natal Museum 20:479-782. [Google Scholar]
- Moore BP. (1974) The larval habits of two species of Sphallomorpha Westwood (Coleoptera: Carabidae, Pseudomorphinae). Journal of the Australian Entomological Society 13:179-183. doi: 10.1111/j.1440-6055.1974.tb02171.x [DOI] [Google Scholar]
- Moore BP. (1979) Chemical defense in carabids and its bearing on phylogeny. In: Erwin TL, Ball GE, Whitehead DR, Halpern AL (Eds) Carabid Beetles: Their Evolution, Natural History, and Classification. Dr. W. Junk Publishers, The Hague, 193–204.
- Moore W, Di Giulio A. (2006) Description and behavoiur of Goniotropis kuntzeni larvae (Coleoptera: Carabidae: Paussinae: Ozaenini) and a key to genera of Paussini larvae. Zootaxa 1111:1-19. [Google Scholar]
- Mossakowski D. (1979) Evolution of habitat preference illustrated by the phylogeny of Chrysocarabus species (Coleoptera, Carabidae). In: Den Boer PJ, Thiele HU, Weber F (Eds) On the Evolution of Behaviour in Carabid Beetles. Miscellaneous Papers, 18, Landbouwhogeschool Wageningen, 103–112.
- Mossakowski D. (2002) The mandible of Maoripamborus: an indication of convergent evolution in Cychrini and Pamborini. In: Szyszko J, Den Boer PJ, Bauer T (Eds) How to protect or what we know about carabid beetles. Warsaw Agricultural University Press, Warsaw, 95–100.
- Mossakowski D. (2005) Morphological or molecular systematics? A case study of Carabus. In: Lövei G, Toft S (Eds) European Carabidology 2003 (Proceedings of the 11th European Carabidologists' Meeting). DIAS Report Plant Production, No. 114, Danish Institute of Agricultural Sciences, 231–241.
- Mossakowski D, Roschen A, Vaje S. (1986) Hybridization in Chrysocarabus. In: Den Boer P, Luff ML, Mossakowski D, Weber F (Eds) Carabid Beetles, their adaptations and dynamics. Gustav Fischer, Stuttgart, 281–295.
- Mossakowski D, Stier J. (1983) Vergleichende Untersuchungen zur Laufgeschwindigkeit der Carabiden. In: Brandmayr P, Den Boer PJ, Weber F (Eds) Report of the fourth meeting of European Carabidologists: the synthesis of field study and laboratory experiment. Centre for Agricultural Publishing and Documentation, Wageningen, The Netherlands, 19–33.
- Muilwijk J, Felix R. (2010) Carabidae - loopkevers. In: Vorst O (Ed) Catalogus van de Nederlandse kevers. Monografieen van de Nederlandse Entomologische Vereniging 11:40-52. [Google Scholar]
- Mullen K, O’Halloran J, Breen J, Giller P, Pithon J, Kelly T. (2008) Distribution and composition of carabid beetle (Coleoptera, Carabidae) communities across the plantation forest cycle- Implications for management. Forest Ecology and Management 256:624-632. doi: 10.1016/j.foreco.2008.05.005 [DOI] [Google Scholar]
- Müller HJ. (1970) Formen der Dormanz bei Insekten. Nova Acta Leopoldina 35:7-25. [Google Scholar]
- Müller-Motzfeld G G. (Ed) (2004) Adephaga 1: Carabidae (Laufkäfer). In: Freude H, Harde KW, Lohse GA, Klausnitzer B (Eds), Die Käfer Mitteleuropas. Vol. 2. Second edition. Elsevier-Spektrum, Akademischer Verlag, München, 521 pp.
- Nabe-Nielsen J, Sibly R, Forchhammer MC, Forbes VE, Topping CJ. (2010) The Effects of Landscape Modifications on the Long-Term Persistence of Animal Populations. PLoS One 5:e8932. doi: 10.1371/journal.pone.0008932 [DOI] [PMC free article] [PubMed]
- Nagel P. (1979a) The classification of Carabidae. In: Den Boer P, Luff ML, Mossakowski D, Weber F (Eds) On the Evolution of Behaviour in Carabid Beetles. Miscellaneous Papers, 18, Landbouwhogeschool Wageningen, 7–14.
- Nagel P. (1979b) Aspects of the evolution of myrmecophilous adaptations in Paussinae (Coleoptera, Carabidae). In: Den Boer P, Luff ML, Mossakowski D, Weber F (Eds) On the Evolution of Behaviour in Carabid Beetles. Miscellaneous Papers, 18, Landbouwhogeschool Wageningen, 15–34.
- Nakagawa S, Cuthill IC. (2007) Effect size, confidence interval and statistical significance: a practical guide for biologists. Biological Reviews 82:591-605. doi: 10.1111/j.1469-185X.2007.00027.x [DOI] [PubMed] [Google Scholar]
- Neculiseanu ZZ, Matalin AV. (2000) A Catalogue of the ground-beetles of the replublic Moldova (Insecta, Coleoptera: Carabidae. Pensoft, Sofia-Moscow, 164 pp.
- Negro M, Casale A, Migliore L, Palestrini C, Rolando A. (2008) Habitat use and movement patterns in the endangered ground beetle species, Carabus olympiae (Coleoptera: Carabidae). European Journal of Entomology 105:105-112. [Google Scholar]
- Nelemans MNE. (1987) Possibilities for flight in the carabid beetle Nebria brevicollis (F.). The importance of food during larval growth. Oecologia 72:502-509. doi: 10.1007/BF00378974 [DOI] [PubMed] [Google Scholar]
- Nettmann H-K. (1986) Carabid systematics and chromosome numbers. In: Den Boer P, Luff ML, Mossakowski D, Weber F (Eds) Carabid Beetles, their adaptations and dynamic. Gustav Fischer, Stuttgart, 235–254.
- Neudecker C, Thiele HU. (1974) Die jahreszeitliche Synchronisation der Gonadenreifung bei Agonum assimile Paxk. (Coleoptera, Carabidae) durch Temperatur und Photoperiode. Oecologia 17:141-157. doi: 10.1007/BF00346277 [DOI] [PubMed] [Google Scholar]
- New TR. (1995) An introduction to invertebrate conservation biology. Oxford University Press, Oxford UK, 208 pp.
- New TR. (2010) Beetles in conservation. Wiley-Blackwell, Chistester, 237 pp. doi: 10.1002/9781444318623 [DOI]
- Nicholson AJ. (1958) Dynamics of insect populations. Annual Review of Entomology 3 107–136. doi: 10.1146/annurev.en.03.010158.000543 [DOI]
- Niehues F-J, Hockmann P, Weber F. (1996) Genetics and dynamics of a Carabus auronitens metapopulation in the Westphalian Lowlands (Coleoptera, Carabidae). Annales Zoologici Fennici 33:85-96. [Google Scholar]
- Niemelä J. (1993) Interspecific competition in ground-beetle assemblages (Carabidae): What have we learned? Oikos 66: 325–335. doi: 10.2307/3544821 [DOI]
- Niemelä J. (1996a) From systematics to conservation – carabidologists do it all. Annales Zoologici Fennici 33:1-4. [Google Scholar]
- Niemelä J. (Ed) (1996b) Population biology and conservation of carabid beetles. Proceedings of the 3rd International Symposium of Carabidology, Kauniainen, Finland. Annales Zoologici Fennici 33:1-241. [Google Scholar]
- Niemelä J. (2001) Carabid beetles (Coleoptera: Carabidae) and habitat fragmentation: a review. European Journal of Entomology 98:127-132. [Google Scholar]
- Niemelä J, Haila Y, Halme E, Pajunen T, Punttila P. (1992) Small-scale heterogeneity in the spatial distribution of carabid beetles in the southern Finnish taiga. Journal of Biogeography 19:173-181. doi: 10.2307/2845503 [DOI] [Google Scholar]
- Niemelä J, Haila Y, Punttila P. (1996) The importance of small-scale heterogeneity in boreal forests: variation in diversity in forest-floor invertebrates across the succession gradient. Ecography 19:352-368. [Google Scholar]
- Niemelä J, Haila Y, Ranta E. (1986) Spatial heterogeneity of carabid beetle dispersion in uniform forests on the Åland Islands, SW Finland. Annales Zoologici Fennici 23:289-296. [Google Scholar]
- Niemelä J, Koivula M, Kotze DJ. (2007) The effects of forestry on carabid beetles (Coleoptera: Carabidae) in boreal forests. Journal of Insect Conservation 11:5-18. doi: 10.1007/s10841-006-9014-0 [DOI] [Google Scholar]
- Niemelä J, Kotze DJ. (2009) Carabid beetle assemblages along urban to rural gradients: A review. Landscape and Urban Planning 92:65-71. doi: 10.1016/j.landurbplan.2009.05.016 [DOI] [Google Scholar]
- Niemelä J, Kotze J, Ashworth A, Brandmayr P, Desender K, New T, Penev L, Samways M, Spence J. (2000) The search for common anthropogenic impacts on biodiversity: a global network. Journal of Insect Conservation 4:3-9. doi: 10.1023/A:1009655127440 [DOI] [Google Scholar]
- Niemelä J, Kotze DJ, Venn S, Penev L, Stoyanov I, Spence J, Hartley D, Montes de Oca E. (2002) Carabid beetle assemblages (Coleoptera, Carabidae) across urban-rural gradients: an international comparison. Landscape Ecology 17:387-401. doi: 10.1023/A:1021270121630 [DOI] [Google Scholar]
- Niemelä J, Ranta E, Haila Y. (1985) Carabid beetles in lush forests patches on the Åland Islands, south-west Finland: an island-mainland comparison. Journal of Biogeography 12:109-120. doi: 10.2307/2844835 [DOI] [Google Scholar]
- Niemelä J, Spence J. (1991) Distribution and abundance of an exotic ground-beetle (Carabidae): a test of community impact. Oikos 62:351-359. doi: 10.2307/3545500 [DOI] [Google Scholar]
- Noonan GR. (1985) The influence of dispersal, vicariance and refugia on patterns of biogeographical distributions of the beetle family Carabidae. In: Ball GE (Ed) Taxonomy, phylogeny and zoogeography of beetles and ants: a volume dedicated to the memory of Philip Jackson Darlington, Jr (1904–1983). Dr. W. Junk, Dordrecht, Boston, Lancaster: 322–349.
- Noonan GR, Ball GE, Stork NE. (1992) The biogeography of ground beetles of maintains and islands. Intercept, Andover, Hampshire, 256 pp.
- Noordijk J. (2009) Arthropods in linear elements, occurrence, behaviour and conservation management. PhD thesis, University of Wageningen, Wageningen, the Netherlands, 188 pp.
- Noordijk J, Prins D, de Jonge M, Vermeulen R. (2006) Impact of a road on the movements of two ground beetle species (Coleoptera: Carabidae). Entomologica Fennica 17:276-283. [Google Scholar]
- Noordijk J, Vermeulen R. (2009) Loopkevers in Drenthe. Stichting Willem Beyerinck Biologisch Station, Loon, 34 pp.
- Norris MJ. (1959) The influence of day-length on imaginal diapause in the red locust, Nomadacris septemfasciata (Serv.). Entomologia Experimentalis et Applicata 2:154-168. doi: 10.1111/j.1570-7458.1959.tb02105.x [DOI] [Google Scholar]
- Norris MJ. (1965) The influence of constant and changing photoperiods on imaginal diapause in the red locust (Nomadacris septemfasciata Serv.). Journal of Insect Physiology 11:1105-1119. doi: 10.1016/0022-1910(65)90181-2 [DOI] [Google Scholar]
- Ober KA, Maddison DR. (2008) Phylogenetic relationships of tribes within Harpalinae (Coleoptera: Carabidae) as inferred from 28S ribosomal DNA and the wingless gene. Journal of Insect Science 8:63, 32 pp. http://www.insectscience.org/8.63/ [DOI] [PMC free article] [PubMed]
- Obydov D. (2002) Révision du genre Callisthenes. Magellanes, Andrésy, 125 pp.
- O’Hara RB, Kotze DJ. (2010) Do not log-transform count data. Methods in Ecology and Evolution 1:118-122. doi: 10.1111/j.2041-210X.2010.00021.x [DOI] [Google Scholar]
- Ortuño VM, Toribio M. (2005) Carabidae de la Península Ibérica y Baleares – Vol. 1. Trechinae, Bembidiini. Argania edition, Barcelona, 455 pp.
- Ouborg NJ, Vriezen WH. (2007) An ecologist's guide to ecogenomics. Journal of Ecology 95:8-16. doi: 10.1111/j.1365-2745.2006.01197.x [DOI] [Google Scholar]
- Paarmann W. (1970) Untersuchungen über die Jahresrhythmik von Laufkäfern (Coleoptera, Carabidae) in der Cyrenaika (Libyen, Nordafrika). Oecologia 5:325-333. doi: 10.1007/BF00815498 [DOI] [PubMed] [Google Scholar]
- Paarmann W. (1973) Bedeutung der Larvenstadien für die Fortpflanzungsrhythmik der Laufkäfer Broscus laevigatus Dej. und Orthomus atlanticus Fairm. (Col., Carab.) aus Nordafrika. Oecologia 13:81-92. doi: 10.1007/BF00379621 [DOI] [PubMed] [Google Scholar]
- Paarmann W. (1974) Der Einfluß von Temperatur und Lichtwechsel auf die Gonadenreifung des Laufkäfers Broscus laevigatus Dej. (Col., Carab.) aus Nordafrika. Oecologia 15:87-92. doi: 10.1007/BF00345230 [DOI] [PubMed] [Google Scholar]
- Paarmann W. (1975) Freilanduntersuchungen in Marokko (Nordafrika) zur Jahresrhythmik von Carabiden (Col., Carab.) und zum Mikroklima im Lebensraum der Käfer. Zoologische Jahrbücher. Abteilung Systematik 102:72-88. [Google Scholar]
- Paarmann W. (1976a) Die Jahresrhythmik und ihre Steuerung bei nord- und zentralafrikanischen Laufkäfern (Col., Carab.). Habiltitationsschrift, Universität Würzburg, 139 pp.
- Paarmann W. (1976b) Jahreszeitliche Aktivität und Fortpflanzungsrhythmik von Laufkäfern (Col., Carab.) im Kivugebiet (Ost-Zaire, Zentralafrika). Zoologische Jahrbücher Abteilung Systematik 103:311-354. [Google Scholar]
- Paarmann W. (1976c) Die Bedeutung exogener Faktoren für die Gonadenreifung von Orthomus barbarus atlanticus Fairm. (Col., Carab.) aus Nordafrika. Entomologia Experimentalis et Applicata 19:23-36. doi: 10.1111/j.1570-7458.1976.tb02578.x [DOI] [Google Scholar]
- Paarmann W. (1976d) The annual periodicity of the polyvoltine ground beetle Pogonus chalceus Marsh. (Col., Carabidae) and its control by environmental factors. Zoologischer Anzeiger 196:150-160. [Google Scholar]
- Paarmann W. (1979) Ideas about the evolution of the various annual reproduction rhythms in carabid beetles of the different climatic zones. Miscellaneous Papers: 18, Landbouw Hogeschool Wageningen, 119–132.
- Paarmann W. (1985) Larvae preying on ant broods: an adaptation of the desert carabid beetle Graphipterus serrator Forskal (Col., Carabidae) to arid environments. Journal of Arid Environments 9:210-214. [Google Scholar]
- Paarmann W. (1986) Seasonality and its control by environmental factors in tropical ground beetles (Col., Carabidae). In: Den Boer PJ, Luff ML, Mossakowski D, Weber F (Eds) Carabid Beetles Their Adaptations and Dynamics. Gustav Fischer Verlag, Stuttgart-New York, 157–171.
- Paarmann W. (1990)Poecilus lepidus Leske (Carabidae, Coleoptera), a species with the ability to be a spring and autumn breeder. In: Stork NE (Ed) The Role of Ground Beetles in Ecological and Environmental Studies. Intercept, Andover, Hampshire, 259–267.
- Paarmann W. (1994) Temperature and photoperiodic influence on developmental stages and adults of the subarctic carabid beetle Pterostichus adstrictus Eschscholtz. In: Desender K, Dufrêne M, Loreau M, Luff ML, Maelfait J-P (Eds) Carabid Beetles: Ecology and Evolution, Kluwer Academic Publishers, Dordrecht 201–205.
- Paarmann W. (2002) Determination of food quality for the successful development of seed-feeding ground beetles (Col., Carabidae) from tropical lowland rain forests. In: Szyszko J, den Boer PJ, Bauer T (Eds) How to Protect or What We Know About Carabid Beetles. Warszaw Agricultural University Press, Warszaw, 45–57.
- Paarmann W, Adis J, Stork N, Gutzmann B, Stumpe P, Staritz B, Bolte H, Kuppers S, Holzkamp K, Niers C, da Fonseca CRV. (2001) The structure of ground beetle assemblages (Coleoptera : Carabidae) at fig fruit falls (Moraceae) in a terra firme rain forest near Manaus (Brazil). Journal of Tropical Ecology 17:549-561. doi: 10.1017/S0266467401001419 [DOI] [Google Scholar]
- Paarmann W, Erbeling L, Spinnler K. (1986) Ant and ant brood preying larvae: An adaptation of carabid beetles to arid environments. In: Den Boer PJ, Luff ML, Mossakowski D, Weber F (Eds) Carabid Beetles Their Adaptations and Dynamics. Gustav Fischer Verlag, Stuttgart-New York, 79–90.
- Paarmann W, Faust N, Arndt E, Luchtrath I, Rohe W. (2006) Constant seed size and mandible growth - a fundamental problem for granivorous ground beetle larvae (Coleoptera : Carabidae). Entomologica Fennica 17:334-339. [Google Scholar]
- Paarmann W, Paarmann D. (1997) Studies on the biology of a canopy-dwelling carabid beetle collected by canopy fogging in the rainforest of Sulawesi (Indonesia). In: Stork NE, Adis J, Didham RK (Eds) Canopy Arthropods. Chapman & Hall, London, England, 433–441.
- Paarmann W, Stork N. (1987) Seasonality of ground beetles (Coleoptera: Carabidae) in the rain forest of N. Sulawesi (Indonesia). Insect Science and its Application 8:483-487. [Google Scholar]
- Palmén E. (1944) Die anemohydrochore Ausbreitung der Insekten als zoogeographischer Faktor. Annales Zoologici Societatis Zoologicae Botanicae Fennicae Vanamo, Helsiki, 262 pp.
- Panzer R, Schwartz MW. (1998) Effectiveness of a vegetation-based approach to insect conservation. Conservation Biology 12:693-702. doi: 10.1046/j.1523-1739.1998.97051.x [DOI] [Google Scholar]
- Parry ML, Canziani OF, Palutikof JP, van der Linden PJ, Hanson CE.Eds (2007) Climate change 2007: impacts, adaptation and vulnerability. Cambridge University Press, Cambridge, 93 pp.
- Pawson SM, Brockerhoff EG, Meenken ED, Didham RK. (2008) Non-native plantation forests as alternative habitat for native forest beetles in a heavily modified landscape. Biodiversity and Conservation 17:1127-1148. doi: 10.1007/s10531-008-9363-y [DOI] [Google Scholar]
- Pearson DL, Vogler AP. (2001) Tiger beetles – The evolution, ecology and diversity of the cicindelids. Comstock Publishing Associates. Cornell University Press, Ithaca, London, 333 pp.
- Pedersen JC, Wind P. (2009) Den danske Rødliste. Danmarks miljoundersogelser, National Environmental Research Institute. URL: http://www2.dmu.dk/1_Om_DMU/2_Tvaer-funk/3_fdc_bio/projekter/redlist/artsgrupper.asp [Access date 15 June 2009]
- Penev L, Erwin T, Assmann T.Eds (2008) Back to the roots and back to the future - Towards a new synthesis between taxonomic, ecological and biogeographical approaches in carabidology. Proceedings of the 13th European Carabidologists Meeting. Pensoft, Sofia-Moscow, 509 pp.
- Perez-Zaballos JM. (1985) Paralelismo fenologico en Brachinus variventris Schaufuss, 1862 y Anchomenus dorsalis (Pontoppidan, 1863). (Coleoptera, Carabidae). Actas do Congresso Iberico de Entomologia 2:85-92. [Google Scholar]
- Persigehl M, Lehmann S, Vermeulen HJW, Rosenkranz B, Falke B, Assmann T. (2004) Kolonisation restituierter Sandrasen im Darmstädter Flugsandgebiet und im mittleren Emsland durch Laufkäfer. NNA-Berichte 1/2004 161–177.
- Petit S. (1994) Diffusion of forest carabid beetles in hedgerow network landscapes. In: Desender K, Dufrêne M, Loreau M, Luff ML, Maelfait J-P (Eds) Carabid beetles: ecology and evolution. Series Entomologica 51:337-341. [Google Scholar]
- Petit S, Burel F. (1998) Effects of landscape dynamics on the metapopulation of a ground beetle (Coleoptera, Carabidae) in a hedgerow network. Agriculture, Ecosystems & Environment 69:243-252. doi: 10.1016/S0167-8809(98)00111-X [DOI] [Google Scholar]
- Petit S, Usher MB. (1998) Biodiversity in agricultural landscapes: the ground beetle communities of woody uncultivated habitats. Biodiversity and Conservation 7:1549-1561. doi: 10.1016/j.ecolmodel.2005.07.027 [DOI] [Google Scholar]
- Pichancourt JB, Burel F, Auger P. (2006) Assessing the effect of habitat fragmentation on population dynamics: An implicit modelling approach. Ecological Modelling 192:543-556. [Google Scholar]
- Pizzolotto R. (2009) 30 years of carabid sampling in Italy: a data bank for studying local climate change. In: Hengeveld R, Noordijk J, Opsteeg T, Turin H, Vermeulen HJW (Eds) Abstracts of the XIV European Carabidologists Meeting, Westerbork, the Netherlands, 24.
- Plachter H. (1986) Composition of the carabid beetle fauna of natural riverbanks and of man-made secondary habitats. In: Den Boer PJ, Thiele, H-U, Weber F (Eds) Carabid beetles, their adaptations and dynamics. Gustav Fischer, Stuttgart, New York, 509–535.
- Plat S, Kuivenhoven P and Van Dijk T. (1995) Hedgerows: suitable corridors for ground-dwelling forest carabid beetles? Proceedings of the section Experimental and Applied Entomology of the Netherlands Entomological Society (N.E.V. ) 6:73-75. [Google Scholar]
- Porta A (1923–1959) Fauna Coleopterorum Italica Vol. I. Adephaga. Piacenza. (1923): 1–285, Supplementum I-III. San Remo (1934, 1949, 1959): 1–208, 1–386, 1–344.
- Prins D, Van Vliet A, Vermeulen R. (2007) Invloed van klimaatverandering op de fenologie en populatiegrootte van loopkevers, een onderzoek op basis van de langstlopende continue meetreeks aan loopkevers ter wereld. Rapport Stichting WBBS, Loon, 82 pp.
- Prüser F, Brückner M, Mossakowski D. (2000) Colonisation of Canary Islands by Carabus species: evidence from different character complexes. In: Brandmayr P, Lövei GL, Zetto Brandmayr T, Casale A, Vigna Taglianti A (Eds) Natural history and applied ecology of carabid beetles. Pensoft, Sofia, 45–52.
- Prüser F, Mossakowski D. (1998) Conflicts in phylogenetic relationships and dispersal history of the supertribe Carabitae (Coleoptera: Carabidae). In: Ball GE, Casale A, Vigna Taglianti A (Eds) Phylogeny and classification of Caraboidea (Coleoptera: Adephaga). Museo Regionale di Scienze Naturali, Torino, 297–328.
- Purtauf T, Dauber J, Wolters V. (2004) Carabid communities in the spatio-temporal mosaic of a rural landscape. Landscape and Urban Planning 67:185-193. doi: 10.1016/S0169-2046(03)00038-0 [DOI] [Google Scholar]
- Purtauf T, Dauber J, Wolters V. (2005) The response of carabids to landscape simplification differs between trophic groups. Oecologia 142:458-464. doi: 10.1007/s00442-004-1740-y [DOI] [PubMed] [Google Scholar]
- Putchkov A. (2011) The ground beetles of Ukraine(Coleoptera, Carabidae). In: Kotze DJ, Assmann T, Noordijk J, Turin H, Vermeulen R (Eds) Carabid Beetles as Bioindicators: Biogeographical, Ecological and Environmental Studies. ZooKeys 100:503-516. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Rainio J, Niemelä J. (2003) Ground beetles (Coleoptera: Carabidae) as bioindicators. Biodiversity and Conservation 12:487-506. doi: 10.1023/A:1022412617568 [DOI] [Google Scholar]
- Ranta E, Ås S. (1982) Non random colonization of habitat islands by carabid beetles. Annales Zoologici Fennici 19:175-181. [Google Scholar]
- Rasplus, J-Y, Meusnier S, Mondor G, Piry S, Cornuet J-M. (2000) Microsatellite analysis of genetic population structure in the endangered beetle: Carabus solieri (Carabidae). In: Brandmayr P. Lövei GL, Zetto Brandmayr T, Casale A, Vigna Taglianti A (Eds) Natural history and applied ecology of carabid beetles. Pensoft, Sofia, 11–24.
- Rassi P, Alanen A, Kanerva T, Mannerkoski I.Eds (2001) Suomen lajien uhanalaisuus 2000. Finnish Ministry of Environment and Finnish Environment Centre, Helsinki, 432 pp.
- Reitter E. (1908) Fauna Germanica. Die Käfer des Deutschen Reiches I Band. K.G. Lutz Verlag, Stuttgart: 248 pp, 40 plates.
- Ribera I, Montagud S, Teruel S, Bellés X. (2006) Molecular data support the inclusion of Ildobates neboti Español in Zuphiini (Coleoptera: Carabidae: Harpalinae). Entomologica Fennica 17:207-213. [Google Scholar]
- Richard E, Gosselin F, Lhonoré J. (2004). Short-term and mid-term response of ground beetle communities (Coleoptera: Carabidae) to disturbance by regeneration felling. In: Honnay O, Bossuyt B, Verheyen K, Hermy M (Eds) Forest Biodiversity: Lessons from history for conservation. Oxon, UK, IUFRO Res. Ser. CAB International, 179–192. doi: 10.1079/9780851998022.0179 [DOI]
- Riecken U, Raths U. (1996) Use of radio telemetry for studying dispersal and habitat use of Carabus coriaceus L. Annales Zoologici Fennici 33:109-116. [Google Scholar]
- Roff D. (1986) The evolution of wing dimorphism in insects. Evolution40: 1009–1020. doi: 10.2307/2408759 [DOI] [PubMed]
- Roig-Juñent S. (1998) Cladistic relationships of the tribe Broscini (Coleoptera: Carabidae). In: Ball GE, Casale A, Vigna Taglianti A (Eds) Phylogeny and classification of Caraboidea (Coleoptera: Adephaga). Museo Regionale di Scienze Naturali, Torino, 343–358.
- Ruiz C, Serrano J. (2006) Molecular phylogeny and systematics of Calathus Bonelli (Coleoptera: Carabidae: Sphodrini) based on mitochondrial cox1-cox2 sequences. Entomologica Fennica 17:214-220. [Google Scholar]
- Rushton SP, Eyre MD, Luff ML. (1990) The effects of scrub management on the ground beetles of oolitic limestone grassland at Castor Hanglands national nature reserve, Cambridgeshire, UK. Biological Conservation 51:97-111. doi: 10.1016/0006-3207(90)90105-X [DOI] [Google Scholar]
- Saarikivi J, Idström L, Venn S, Niemelä J, Kotze DJ. (2010) Carabid beetle assemblages associated with urban golf courses in the greater Helsinki area. EuropeanJournal of Entomology 107:553-561. [Google Scholar]
- Sadler JP, Small EC, Fiszpan H, Telfer MG, Niemelä J. (2006) Investigating environmental variation and landscape characteristics of an urban-rural gradient using woodland carabid assemblages. Journal of Biogeography 33:1126-1138. doi: 10.1111/j.1365-2699.2006.01476.x [DOI] [Google Scholar]
- Sætersdal M, Gjerde I, Blom HH. (2005) Indicator species and the problem of spatial inconsistency in nestedness patterns. Biological Conservation 122:305-316. doi: 10.1016/j.biocon.2004.07.020 [DOI] [Google Scholar]
- Sahlberg J. (1868) Bidrag till kännedomen om Finlands Dimorpha insekt-arter (Engl. Contributions to the knowledge of Finland’s Dimorpha species) Notiser ur Sällskapets pro Fauna et Flora Fennica förhandlingar9 (1867) Helsinki.
- Salt G. (1928). Notes on the life history of Pelecium sulcatum Guerin. Psyche 35:131-134. [Google Scholar]
- Samways MJ. (2005) Insect diversity conservation. Cambridge University Press, Cambridge, 250 pp. doi: 10.1017/CBO9780511614163 [DOI]
- Samways MJ. (2007) Insect conservation: a synthetic management approach. Annual Review of Entomology 52:465-487. doi: 10.1146/annurev.ento.52.110405.091317 [DOI] [PubMed] [Google Scholar]
- Sander A, Purtauf T, Wolters V and Dauber J. (2006) Landscape genetics of the widespread ground-beetle Carabus auratus in an agricultural region. Basic and Applied Ecology 7:555-564. doi: 10.1016/j.baae.2006.01.001 [DOI] [Google Scholar]
- Sapia M, Lövei GL, Elek Z. (2006) Effects of varying sampling effort on the observed diversity of carabid (Coleoptera: Carabidae) assemblages in the Danglobe Project, Denmark. Entomologica Fennica 17:345-350. [Google Scholar]
- Sasakawa K. (2007) Laboratory studies on larval feeding habits of Amara macronota (Coleoptera: Carabidae: Zabrini). Applied Entomology and Zoology 42:669-674. doi: 10.1303/aez.2007.669 [DOI] [Google Scholar]
- Sasakawa K. (2009) Effects of maternal diet on fecundity and larval development in the 'primitive' granivorous carabid Amara (Curtonotus) macronota. Entomologia Experimentalis et Applicata 130:106-112. doi: 10.1111/j.1570-7458.2008.00799.x [DOI] [Google Scholar]
- Saska P. (2005) Contrary food requirements of the larvae of two Curtonotus (Coleoptera: Carabidae: Amara) species. Annals of Applied Biology 147:139-144. doi: 10.1111/j.1744-7348.2005.00016.x [DOI] [Google Scholar]
- Saska P. (2008) Effect of diet on the fecundity of three carabid beetles. Physiological Entomology 33:188-192. doi: 10.1111/j.1365-3032.2008.00618.x [DOI] [Google Scholar]
- Saska P, Honek A. (2004) Development of the beetle parasitoids, Brachinus explodens and B. crepitans (Coleoptera: Carabidae). Journal of Zoology, London 262:29-36. doi: 10.1017/S0952836903004412 [DOI] [Google Scholar]
- Saska P, Honek A. (2005) Development of the ground-beetle parasitoids, Brachinus explodens and B. crepitans (Coleoptera: Carabidae): effect of temperature. In: Lövei GL, Toft S (Eds) European Carabidology 2003. Proceedings of the 11th European Carabidologists’ Meeting. DIAS Report 114 :267-274..
- Saska P, Honek A. (2008) Synchronization of a coleopteran parasitoid, Brachinus spp. (Coleoptera: Carabidae), and its host. Annals of the Entomological Society of America 101:533-538. doi: 10.1603/0013-8746(2008)101[533:SOACPB]2.0.CO;2 [DOI] [Google Scholar]
- Saska P, Jarošík V. (2001) Laboratory study of larval food requirements in nine species of Amara (Coleoptera: Carabidae). Plant Protection Science 37:103-110. [Google Scholar]
- Saska P, Martinkova Z, Honek A. (2010) Temperature and rate of seed consumption by ground beetles (Carabidae). Biological Control 52:91-95. doi: 10.1016/j.biocontrol.2009.07.016 [DOI] [Google Scholar]
- Saska P, Vodde M, Heijerman T, Westerman P, van der Werf W. (2007) The significance of a grassy field boundary for the spatial distribution of carabids within two cereal fields. Agriculture, Ecosystems and Environment 122:427-434. doi: 10.1016/j.agee.2007.02.013 [DOI] [Google Scholar]
- Schjøtz-Christensen B. (1965) Biology and population studies of Carabidae of the Corynephoretum. Natura Jutlandica 11: 1–173, tables 1–89.
- Schjøtz-Christensen B. (1966) Biology of some ground beetles (Harpalus Latr.) of the Corynephoretum. Natura Jutlandica 12:225-229. [Google Scholar]
- Schjøtz-Christensen B. (1968) Some notes on the biology of Carabus hortensis L. (Coleoptera, Carabidae). Natura Jutlandica 14:127-154. [Google Scholar]
- Schremmer F. (1960) Beitrag zur Biologie von Ditomus clypeatus Rossi, eines körnersammelnden Carabiden. Zeitschrift der Arbeitsgemeinschaft Östereichischer Entomologen 3:140-146. [Google Scholar]
- Schüle P, Terlutter H. (1998) Rote Liste der gefährdeten Sandlaufkäfer und Laufkäfer (Coleoptera: Cicindelidae, Carabidae) in Nordrhein-Westfalen. Ergänzter Nachdruck aus Angewandte Carabidologie 1:51-62. [Google Scholar]
- Schweiger O, Maelfait JP, Van Wingerden W, Hendrickx F, Billeter R, Speelman, M, Augenstein I, Aukema B, Aviron S, Bailey D, Bukacek R, Burel F, Diekotter T, Dirksen J, Frenzel M, Herzog F, Liira J, Roubalova M. (2005) Quantifying the impact of environmental factors on arthropod communities in agricultural landscapes across organizational levels and spatial scales. Journal of Applied Ecology 42:1129-1139. doi: 10.1111/j.1365-2664.2005.01085.x [DOI] [Google Scholar]
- Schwerk A. (2000) Ecological aspects of carabid beetles coenoses (Coleoptera: Carabidae) on industrial fallow grounds in the Ruhr Valley area. In: Brandmayr P, Lövei G, Zetto Brandmayr T, Casale A, Taglianti V (Eds) Natural history and applied ecology of carabid beetles, Pensoft, 277–287.
- Schwerk A. (2008) Model of the rate of succession of epigeic carabid beetles (Coleoptera: Carabidae) on degraded areas. Instytut Badawczy Leśnictwa, Sękocin Stary, 71 pp.
- Sciaky R. (1987) Revisione delle specie paleartiche occidental del genere Ophonus Dejean. Memorie della Società Entomologia Italiana 65:29-120. [Google Scholar]
- Serrano J. (1986) A karyotypical approach to carabid evolution (Coleoptera). In: Den Boer P, Luff ML, Mossakowski D, Weber F (Eds) Carabid Beetles, their Adaptations and Dynamics. Gustav Fischer, Stuttgart, 221–234.
- Serrano J. (1992) Geographic isolation and karyotypic evolution of carabid beetles (Coleoptera: Carabidae). In: Noonan GR, Ball GE, Stork NE (Eds) The biogeography of ground beetles (Coleoptera: Carabidae) from mountains and islands. Intercept Publ., Andover, 235–243.
- Serrano J. (2003) Catálogo de los Carabidae (Coleoptera) de la Península Ibérica. Monografías de la Sociedad Entomológica Aragonesa, Zaragoza, 130 pp.
- Serrano J, Galián J. (1998) A review of karyotypic evolution and phylogeny of carabid beetles (Coleoptera). In: Ball GE, Casale A, Vigna Taglianti A (Eds) Phylogeny and classification of Caraboidea (Coleoptera: Adephaga). Museo Regionale di Scienze Naturali, Torino, 191–228.
- Serrano J, Galián J, Ortiz AS. (1994) Karyotypic data and current taxonomic ideas of the tribe Harpalini (Coleoptera, Carabidae). In: Desender K, Dufrêne M, Loreau M, Luff ML, Maelfait J-P (Eds) Carabid beetles: Ecology and evolution. Kluwer Academic Press, Dordrecht, 55–61.
- Serrano J, Koivula M, Lövei G.Eds (2006) Proceedings of the 12th European Carabidologists’ Meeting. Entomologica Fennica 17: 35: 0 pp. [Google Scholar]
- Sharova ICH. (1981) Life forms of carabids (Coleoptera, Carabidae), Ghilarov MS (Ed) Nauka Publishers, Moscow, 360 pp.
- Silvestri F. (1904) Contribuzione alla conoscenza della metamorphosi e dei costumi della Lebia scapularis Fourcr. con descrizione dell' aparato sericiparo della larva. Redia 2:68-84. [Google Scholar]
- Similä M, Kouki J, Mönkkönen M, Sippola A-L, Huhta E. (2006) Co-variation and indicators of species diversity: can richness of forest-dwelling species be predicted in northern boreal forests? Ecological Indicators 6: 686–700. doi: 10.1016/j.ecolind.2005.08.028 [DOI]
- Skłodowski JJW. (2006) Anthropogenic transformation of ground beetle assemblages (Coleoptera: Carabidae) in Bialowieza forest, Poland: from primeval forest to managed woodlands of various ages. Entomologica Fennica 17:296-309. [Google Scholar]
- Skłodowski J, Garbalińska P. (2010) Zmiany zgrupowań biegaczowatych w drzewostanach zaburzonych. In: Skłodowski J (Ed) Monitoring zoo indykacyjny regeneracji ekosystemów leśnych Puszczy Piskiej zaburzonych przez huragan. Warsaw Agricultural University Press, Warsaw, 145–189.
- Skoupý V. (2004) Ground-beetles (Coleoptera: Carabidae) of the Czech and Slovak Republics of Jan Pulpán’s collection. Skoupý, Praha, 213 pp.
- Sokal RR, Rohlf FJ. (1995)Biometry. 3rd edition. Freeman and Company, New York, 887 pp.
- Stork NE. (Ed) (1990) The role of ground beetles in ecological and environmental studies. Proceedings of the 7th European Carabidologists’ Meeting. Intercept, Andover, Hampshire, 424 pp.
- Stork N, Paarmann W. (1992) Reproductive seasonality of the ground and tiger beetle (Coleoptera: Carabidae, Cicindelidae) fauna in N. Sulawesi, Indonesia. Studies on Neotropical Fauna and Environment 27:101-115. doi: 10.1080/01650529209360871 [DOI] [Google Scholar]
- Strand A. (1970) Additions and corrections to the Norwegian part of the Catalogus Coleopterorum Fennoscandiae et Daniae (Lindroth, 1960). Norsk entomologisk Tidsskrift 17:125-126. [Google Scholar]
- Šustek Z. (1987) Changes in body size structure of carabid communities (Coleoptera, Carabidae) along an urbanisation gradient. Biologia (Bratislava)42: 145–156.
- Šustek Z. (1992) Changes in the representation of carabid life forms along an urbanisation gradient (Coleoptera, Carabidae). Biologia (Bratislava) 47:417-430. [Google Scholar]
- Sutherland WJ (Ed) (1998) Conservation science and action. Blackwell Science, Oxford, UK, 376 pp.
- Szyszko J. (1981) The organization of a field station in Niedzwiady for experimental research on carabid beetles. In Brandmayr P, Den Boer PJ, Weber F (Eds) Ecology of carabids: The synthesis of field study and laboratory experiments - Report of the Fourth meeting of European Carabidologists, University of Münster, Westphalia, 183–189.
- Szyszko J. (1986a) The occurrence of Carabidae (Coleoptera) in pine stands in fresh coniferous forest habitats in the district Niedżiady. In: den Boer PJ, Grüm L, Szyszko J (Eds) Feeding behaviour and accessibility of food for carabid beetles. Warsaw Agricultural University Press, Warsaw, 133–147.
- Szyszko J. (1986b) What is the habitat for carabid beetles (Coleoptera, Carabidae)? In: Den Boer PJ, Grüm L, Szyszko J (Eds) Feeding behaviour and accessibility of food for carabid beetles. Warsaw Agricultural University Press, Warsaw, 149–156.
- Szyszko J. (1990) Planning of prophylaxis in threatened pine forest biocoenoses based on an analysis of the fauna of epigeic Carabidae. Warsaw Agricultural University Press, Warsaw, 96 pp.
- Szyszko J. (2004) Foundations of Poland's cultural landscape protection – conservation policy. Dieterich M, van der Straaten J (Eds) Cultural landscapes and land use. Kluwer Academic Publishers, The Netherlands, 95–110.
- Szyszko J. (2010): Formowanie się zgrupowań epigeicznych biegaczowatych w zależności od wieku drzewostanu i historii użytkowania gleb. In: Szyszko J, Tobolski K (Eds) Podstawy Kompensacji Przyrodniczej. Wyższa Szkola Kultury Społecznej i Medialnej w Toruniu Press, Toruń, 126–138.
- Szyszko J, Den Boer PJ, Bauer T.Eds (2002) How to protect or what we know about carabid beetles. Proceedings of the 10th European Carabidologists’ Meeting. Warsaw Agricultural University Press, Warsaw, 1–378.
- Szyszko J, Den Boer PJ, Vermeulen HJR. (1996) Survival and reproduction in relation to habitat quality and food availability for Pterostichus oblongopunctatus F. (Carabidae, Col.). Acta Jutlandica 71:25-40. [Google Scholar]
- Szyszko J, Schwerk A, Malczyk J. (2011) Animals as an indicator of carbon sequestration and valuable landscapes. In: Kotze DJ, Assmann T, Noordijk J, Turin H, Vermeulen R (Eds) Carabid Beetles as Bioindicators: Biogeographical, Ecological and Environmental Studies. ZooKeys 100:564-573. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Szyszko J, Vermeulen HJW, Klimaszewski K, Abs M, Schwerk A. (2000) Mean Individual Biomass (MIB) of ground beetles (Carabidae) as an indicator of the state of the environment. In: Brandmayr P, Lövei G, Zetto Brandmayr T, Casale A, Taglianti AV (Eds) Natural history and applied ecology of carabid beetles. Proceedings of the IX European Carabidologists' Meeting (26–31 July 1998, Camigliatello, Cosenza, Italy). Pensoft, Sofia-Moscow, 289–294.
- Taboada A, Kotze DJ, Salgado JM, Tárrega R. (2006a) The influence of habitat type on the distribution of carabid beetles in traditionally managed "dehesa" ecosystems in NW Spain. Entomologica Fennica 17:284-293. [Google Scholar]
- Taboada A, Kotze DJ, Tárrega R, Salgado JM. (2006b) Traditional forest management: Do carabid beetles respond to human-created vegetation structures in an oak mosaic landscape? Forest Ecology and Management 237: 436–449. doi: 10.1016/j.foreco.2006.09.077 [DOI]
- Taboada A, Kotze DJ, Tárrega R, Salgado JM. (2008) Carabids of differently aged reforested pinewoods and a natural pine forest in a historically modified landscape. Basic and Applied Ecology 9:161-171. doi: 10.1016/j.baae.2007.01.004 [DOI] [Google Scholar]
- Tanner RA, Gange AC. (2005) Effects of golf courses on local biodiversity. Landscape and Urban Planning 71:137-146. doi: 10.1016/S0169-2046(04)00034-9 [DOI] [Google Scholar]
- Taylor RJ, Doran N. (2001) Use of terrestrial invertebrates as indicators of the ecological sustainability of forest management under the Montreal Process. Journal of Insect Conservation 5:221-231. doi: 10.1023/A:1013397410297 [DOI] [Google Scholar]
- Telfer MG, Eversham BC. (1996) Ecology and conservation of heathland Carabidae in eastern England. Annales Zoologici Fennici 33:133-138. [Google Scholar]
- Terlutter H. (1990) An allele gradient of an esterase gene locus as result of recent gene flow: electrophoretic investigations of Carabus auronitens F. (Col. Carabidae). In: Stork NE (Ed) The role of ground beetles in ecological and environmental studies, Intercept, 359–364.
- Thacker JRM. (1996) Carabidologists and fragmented habitats. Trends in Ecology and Evolution 11:103-104. doi: 10.1016/0169-5347(96)81083-4 [DOI] [PubMed] [Google Scholar]
- Thiele H-U. (1971) Wie isoliert sind Populationen von Waldcarabiden in Feldhecken. In: Den Boer PJ (Ed) Dispersal and Dispersal Power of Carabid Beetles. Miscellaneous Papers Landbouwhogeschool Wageningen 8:105-110. [Google Scholar]
- Thiele H-U. (1977) Carabid beetles in their Environment: a Study on Habitat Selection by Adaptations in Physiology and Behaviour. Springer, Berlin, 369 pp.
- Thomas CFG, Holland JM, Brown NJ. (2002) The spatial distribution of carabid beetles in agricultural landscapes. In: Holland JM (Ed) The Agroecology of Carabid Beetles. Intercept Limited, Hampshire, UK, 305–344.
- Thomas CFG, Parkinson L, Marshall EJP. (1998) Isolating the components of activity-density for the carabid beetle Pterostichus melanarius in farmland. Oecologia 116:103-112. doi: 10.1007/s004420050568 [DOI] [PubMed] [Google Scholar]
- Thomas MB. (1990) The role of man-made grassy habitats in enhancing carabid populations in arable land. In: Stork NE (Ed) The role of ground beetles in ecological and environmental studies. Intercept, Andover, Hampshire, 77–85.
- Toft S, Bilde T. (2002) Carabid diets and food value. In: Holland JM (Ed) The Agroecology of Ground Beetles. Intercept, Andover, 81–110.
- Trautner J, Geigenmüller K. (1987) Tiger beetles, ground beetles – Illustrated key to the Cicindelidae and Carabidae of Europe. J. Margraf Verlag, Aichtal, Germany, 488 pp.
- Turin H. (1981) Provisional checklist of the European ground-beetles (Coleoptera, Cicindelidae & Carabidae). Monograph of the Netherlands Entomological Society 9: 24: 9 pp. [Google Scholar]
- Turin H. (2000) De Nederlandse Loopkevers, Verspreiding en Oecologie (Coleoptera: Carabidae). Nederlandse Fauna 3. Nationaal Natuurhistorisch Museum Naturalis, KNNV Uitgeverij & EIS-Nederland, Leiden, 666 pp.
- Turin H, Alders K, Den Boer PJ, Van Essen S, Heijerman, Th, Laane W, Penterman E. (1991) Ecological characterisation of carabid species (Coleoptera, Carabidae) in the Netherlands from thirty years of pitfall sampling. Tijdschrift voor Entomologie 134:279-304. [Google Scholar]
- Turin H, Den Boer PJ. (1988) Changes in the distribution of carabid beetles in the Netherlands since 1880. II. Isolation of habitats and long-term time trends in the occurrence of carabid species with different powers of dispersal (Coleoptera, Carabidae). Biological Conservation 44:179-200. doi: 10.1016/0006-3207(88)90101-2 [DOI] [Google Scholar]
- Turin H, Penev L, Casale A.Eds (2003) The genus Carabus – A Synthesis. Pensoft, Sofia-Moscow, 511 pp.
- Underwood AJ. (2009) Components of design in ecological field experiments. Annales Zoologici Fennici 46:93-111. [Google Scholar]
- Vanbergen AJ, Woodcock BA, Watt AD, Niemelä J. (2005) Effect of land-use heterogeneity on carabid communities at the landscape scale. Ecography 28:3-16. doi: 10.1111/j.0906-7590.2005.03991.x [DOI] [Google Scholar]
- Vanbergen AJ, Woodcock BA, Koivula M, Niemelä J, Kotze DJ, Bolger T, Golden V, Dubs F, Boulanger G, Serrano J, Lencina JL, Serrano A, Aguiar C, Grandchamp A-C, Stofer S, Szél G, Ivits E, Adler P, Markus J, Watt AD. (2010) Trophic level modulates carabid beetle responses to habitat and landscape structure: a pan-European study. Ecological Entomology 35:226-235. doi: 10.1111/j.1365-2311.2010.01175.x [DOI] [Google Scholar]
- Van Dijk ThS, Den Boer PJ. (1992) The life histories and population dynamics of two carabid species on a Dutch heathland. 1. Fecundity and the mortality of immature stages. Oecologia 90:340-352. doi: 10.1007/BF00317690 [DOI] [PubMed] [Google Scholar]
- Van Essen Sj. (1993) Loopkeverinventarisatie Dwingelderveld 1991, SBB, Biologisch Station Wijster, Natuurmonumenten. Grafisch Service Centrum Wageningen. 1–97.
- Van Huizen TH. (1977) The significance of flight activity in the life cycle of Amara plebeja Gyllh. (Coleoptera, Carabidae). Oecologia 29:27-41. doi: 10.1007/BF00345360 [DOI] [PubMed] [Google Scholar]
- Van Huizen TH. (1979) Individual and environmental factors determining flight in carabid beetles. In: Den Boer PJ, Thiele H-U, Weber F (Eds) On the evolution of behaviour in Carabid beetles. Miscellaneous papers, Agricultural University Wageningen 18:199-211. [Google Scholar]
- Venn SJ. (2007) Morphological responses to disturbance in wing-polymorphic carabid species (Coleoptera Carabidae) of managed urban grasslands. Baltic Journal of Coleopterology 7:51-60. [Google Scholar]
- Venn SJ, Kotze DJ, Niemelä J. (2003) Urbanization effects on carabid diversity in boreal forests. European Journal of Entomology 100:73-80. [Google Scholar]
- Ver Hoef JM, Boveng PL. (2007) Quasi-Poisson vs. negative binomial regression: how should we model overdispersed count data? Ecology 88:2766-2772. doi: 10.1890/07-0043.1 [DOI] [PubMed] [Google Scholar]
- Vermeulen HJW. (1986) De invloed van voedselhoeveelheid en voedselkwaliteit op de reproductie, groei en overleving van Pterostichus oblongopunctatus en Calathus erratus. Doctoral thesis, Biological Station, Wijster.
- Vermeulen HJW. (1993) The composition of the carabid fauna on poor sandy road-side verges in relation to comparable open areas. Biodiversity and Conservation 2:331-350. doi: 10.1007/BF00114038 [DOI] [Google Scholar]
- Vermeulen HJW. (1994) Corridor function of a road verge for dispersal of stenotopic heathland ground beetles Carabidae. Biological Conservation 69:339-349. doi: 10.1016/0006-3207(94)90433-2 [DOI] [Google Scholar]
- Vermeulen R, Opsteeg T. (1994) Movements of some carabid beetles in road-side verges. Dispersal in a simulation programme. In: Desender K, Dufrêne M, Loreau M, Luff ML, Maelfait J-P (Eds) Carabid beetles: ecology and evolution. Series Entomologica 51:393-398. [Google Scholar]
- Vermeulen HJW, Opdam PFM. (1995) Effectiveness of roadside verges as dispersal corridors for small ground-dwelling animals: A simulation study. Landscape and Urban Planning 31:233-248. doi: 10.1016/0169-2046(94)01050-I [DOI] [Google Scholar]
- Vermeulen HJW, Opsteeg TJ, Kooij A. (2002) Planning ecological corridors for carabid beetles. The proposed ecological connection in Miud-Drenthe (Netherlands), Terhorsterzand-Scharreveld-Orveltezand. In: Szyszko J, Den Boer PJ, Bauer T (Eds) How to protect or what we know about carabid beetles. Warsaw Agricultural University Press, Warsaw, 133–142.
- Vermeulen R, Spee A. (2005) The Mantingerveld: effects of fragmentation and defragmentation followed by carabid beetles. In: Lövei G, Toft S (Eds) European Carabidology 2003 (Proceedings of the 11th European Carabidologists’ Meeting). DIAS Report Plant Production, No. 114, Danish Institute of Agricultural Sciences, 379–389.
- Vermeulen R, Spee A, Popken R. (2004) Versnippering en ontsnippering in het Mantingerveld; de gevolgen voor Loopkevers. De Levende Natuur 105:67-72. [Google Scholar]
- Vermeulen HJW, Szyszko J. (1992) Cooperation within Europe in a study of Pterostichus oblongopunctatus F. (Coleoptera, Carabidae). The influence of food and habitat quality on the egg production. In: Zombori L, Peregovits L (Eds) Proceedings of the 4th European Congress of Entomology, Hungarian Natural History Museum, Budapest, 592–601.
- Versteirt V, Dekoninck W, Desender K, Grootaert P. (2002) Ground beetle communities as evaluation of reconverted arable land into heathland and dry oligotrophic grasslands (Flanders, Belgium). In: Szyszko J, Den Boer PJ, Bauer T (Eds) How to protect or what we know about carabid beetles, Warsaw Agricultural University Press, 143–154.
- Vigna Taglianti A. (1993) Checklist delle specie della fauna d’Italia, 44 Coleoptera Archostemata, Adephaga 1 (Carabidae). Ministerio dell’Ambiente Comitato Scientifico per la Fauna d’Italia. Edizione Calderini, Bologna, 51 pp.
- Vigna Taglianti A. (2005) Checklist e Corotipi delle specie di Carabidi della fauna Italiana. In: Brandmayr P, Zetto T, Pizzolotto R (Eds) I Coleotteri Carabidi per la valutazione ambientale e la conservazione della biodiversità. APAT, Roma, 186–225.
- Vigna Taglianti A, Santarelli E, Di Giulio A, Oliverio M. (1998) Phylogenetic implications of larval morphology in the tribe Ozaenini (Coleoptera: Carabidae). In: Ball GE, Casale A, Vigna Taglianti A (Eds) Phylogeny and classification of Caraboidea (Coleoptera: Adephaga). Museo Regionale di Scienze Naturali, Torino, 273–295.
- Vinson SB. (1976) Host selection by insect parasitoids. Annual Review of Entomology 21:109-133. doi: 10.1146/annurev.en.21.010176.000545 [DOI] [Google Scholar]
- Vlijm L, van Dijk TS, Wijmans SY. (1968) Ecological studies on carabid beetles. III. Winter mortality in adult Calathus melanocephalus (Linn.). Egg production and locomotory activity of the population which has hibernated. Oecologia 1:304-314. doi: 10.1007/BF00386687 [DOI] [PubMed] [Google Scholar]
- Vogler AP, Barraclough TG. (1998) Reconstructing shifts in diversification rate during the radiation of tiger beetles (Coleoptera). In: Ball GE, Casale A, Vigna Taglianti A (Eds) Phylogeny and classification of Caraboidea (Coleoptera: Adephaga). Museo Regionale di Scienze Naturali, Torino, 251–260.
- Vogler AP, DeSalle R. (1994) Mitochondrial DNA evolution and the application of the phylogenetic species concept in the Cicindela dorsalis complex (Coleoptera: Cicindelidae). In: Desender K, Dufrêne M, Loreau M, Luff ML, Maelfait J-P (Eds) Carabid beetles: Ecology and evolution. Kluwer Acad. Press, Dordrecht, 79–85.
- Vogt S, Kamer N, Mossakowski D. (2005) The phylogeny of African Anthiini beetles (Coleoptera: Carabidae) inferred from mitochondrial NADH dehydrogenase subunit 5 (ND5) DNA sequences. In: Lövei G, Toft S (Eds) European Carabidology 2003 (Proceedings of the 11th European Carabidologists' Meeting). DIAS Report Plant Production, No. 114, Danish Institute of Agricultural Sciences, 391–397.
- Wachmann E, Platen R, Barndt D. (1995) Laufkäfer – Beobachtung, Lebensweise. Naturbuchverlag Augsburg, 293 pp.
- Walter H, Lieth H. (1960) Klimadiagramm Weltatlas, G. Fischer Verlag, Jena, Germany, 253 pp.
- Ward RD. (1979) Metathoracic wing structures as phylogenetic indicators in the Adephaga (Coleoptera). In Erwin TL, Ball GE, Whitehead DR, Halpern AL (Eds) Carabid Beetles: Their Evolution, Natural History, and Classification. Dr. W. Junk Publishers, The Hague, 181–192.
- Weber D, Saska P, Chaboo C. (2008) Carabid beetles (Coleoptera: Carabidae) as parasitoids. In: Capinera JL (Ed) Encyclopedia of Entomology, Springer, New York, Berlin, Heidelberg, Tokyo, 719–721.
- Weber DC, Rowley DL, Greenstone MH, Athanas MM. (2006) Prey preference and host suitability of the predatory and parasitoid carabid beetle, Lebia grandis, for several species of Leptinotarsa beetles. Journal of Insect Science 6 (09), 14 pp. www.insectscience.org/6.09. [DOI] [PMC free article] [PubMed]
- White GC, Bennetts RE. (1996) Analysis of frequency count data using the negative binomial distribution. Ecology 77:2549-2557. doi: 10.2307/2265753 [DOI] [Google Scholar]
- White TCR. (1993) The inadequate environment, Nitrogen and the abundance of animals. Springer-Verlag, Berlin, 425 pp.
- Wickham HF. (1893) Description of the early stages of several North American Coleoptera. Bulletin from the Laboratories of natural history of the State university of Iowa 2:330-332. [Google Scholar]
- Wizen G, Gasith A. (2011) Predation of amphibians by carabid beetles of the genus Epomis found in the central coastal plain of Israel. In: Kotze DJ, Assmann T, Noordijk J, Turin H, Vermeulen R (Eds) Carabid Beetles as Bioindicators: Biogeographical, Ecological and Environmental Studies. ZooKeys 100:181-191. [DOI] [PMC free article] [PubMed] [Google Scholar]
- Woodcock BA, Redhead J, Vanbergen AJ, Hulmes L, Hulmes S, Peyton J, Nowakowski M, Pywell RF, Heard MS. (2010) Impact of habitat type and landscape structure on biomass, species richness and functional diversity of ground beetle. Agriculture, Ecosystems & Environment 139:181-186. doi: 10.1016/j.agee.2010.07.018 [DOI] [Google Scholar]
- Yamada Y, Sasaki H, Harauchi Y. (2010) Effects of narrow roads on the movement of carabid beetles (Coleoptera, Carabidae) in Nopporo Forest Park, Hokkaido. Journal of Insect Conservation 14:151-157. doi: 10.1007/s10841-009-9236-z [DOI] [Google Scholar]
- Zaballos JP, Jeanne C. (1994) Nuevo catalogo de los carábidos (Coleoptera) de la peninsula Iberica. Monografias de la Sociedad Entomológica Aragonesa 1:1-159. [Google Scholar]
- Zalewski M. (2004) Do smaller islands host younger populations? A case study on metapopulations of three carabid species. Journal of Biogeography 31:1139-1148. doi: 10.1111/j.1365-2699.2004.01097.x [DOI] [Google Scholar]
- Zamotajlov A, Sciaky R. (1999) Advances in Carabidology – Papers dedicated to the memory of Prof. Oleg L. Kryzhanovskij. MUISO Publishers, Krasnodar, 472 pp.
- Zetto Brandmayr T. (1976) Nutrizione e allevamento di Carabidi esclucivamente fitofagi: spermofagia larvale di Ophonus ardosiacus Lutsh. Redia 59:197-206. [Google Scholar]
- Zetto Brandmayr T. (1983a) Regolazione esogena dello sviluppo preimmaginale di Ophonus rotundicollis Fairm. e Lab. (ardosiacus Lutsh) (Coleoptera, Carabidae) in dipendenza da temperatura e fotoperiodo. Proceedings XII Congresso Nazionale Italiano Entomologia, Roma, 1980, 105–112.
- Zetto Brandmayr T. (1983b) Life cycle, control and propagation rhythm and fecundity of Ophonus rotundicollis Fairm. et Lab. (Coleoptera, Carabidae, Harpalini) as an adaptation to the main feeding plant Daucus carota L. (Umbelliferae). In: Brandmayr P, Den Boer PJ, Weber F (Eds) Ecology of Carabids: the synthesis of field study and laboratory experiment. ”Haus Rothenberge” Westfalia, 93–103.
- Zetto Brandmayr T. (1990) Spermophagous (seed-eating) ground beetles: first comparison of the diet and ecology of the harpaline genera Harpalus and Ophonus (Col., Carabidae). In: Stork N (Ed) The Role of Ground Beetles in Ecological and Environmental Studies. Intercept, Andover, 307–316.
- Zetto Brandmayr T, Dalpozzo R, De Nino A, De Rose E, Giglio A, Procopio A, Sindona G, Talarico F. (2000a) Siagona europaea Dejean (Coleoptera, Carabidae), an ant predator living with ants: hypothesis about chemical mimicry. Insect Social Life 3:201-208. [Google Scholar]
- Zetto Brandmayr T, De Rose E, Giglio A, Pizzolotto R. (2000b) Morphological and behavioural aspects favouring ant predation in Siagona europaea (Coleoptera, Carabidae). In: Brandmayr P, Lövei GL, Zetto Brandmayr T, Casale A, Vigna-Taglianti A (Eds) Natural History and Applied Ecology of Carabid Beetles. Pensoft, Sofia, 103–109.
- Zetto Brandmayr T, Giglio A, De Rose E. (1998a) Feeding behaviour and food preferences of Siagona europaea Dejean, a myrmecophagous carabid beetle (Coleoptera, Carabidae). Insect Social Life 2:203-207. [Google Scholar]
- Zetto Brandmayr T, Giglio A, Marano I, Brandmayr P. (1998b) Morphofunctional and ecological features in carabid (Coleoptera) larvae. In: Ball GE, Casale A, Vigna Taglianti A (Eds) Phylogeny and Classification of Caraboidea. Museo Regionale di Scienze Naturali, Torino, 449–489.