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Ecology and Conservation Biology of Ground Beetles (Coleoptera: Carabidae) in an Age of Increasing Human Dominance

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Ecology and Conservation Biology of Ground Beetles (Coleoptera: Carabidae) in an Age of Increasing Human Dominance Ecology and conservation biology of ground beetles (Coleoptera: Carabidae) in an age of increasing human dominance Gábor Lövei 2008 3 Table of Contents Introduction......................................................................................................................................5 Part I: The ecology and behaviour of ground beetles…………………...…………………………7 Part II. Methodological innovations……………………………………...………………………23 1. The non-equivalence of the two components of trapping effort: sampling duration and the number of traps…………………………….……………….24 2. Effects of varying sampling regimes on the observed diversity of carabid assemblages …………………………………………......…………………….28 3. Harmonic radar - a method using inexpensive tags to study invertebrate movement on land ..........................................................................................33 Discussion......................................................................................................................................38 Part III. Life history studies…………………………………………………….……..…………43 1. Seasonal activity and reproduction of Anisodactylus signatus , Platynus dorsalis and Brachinus explodens in Hungary …………….…………44 2. Seasonal dynamics and reproductive phenology of ground beetles in fragments of native forest in the Manawatu, North Island, New Zealand .....................54 Discussion......................................................................................................................................63 Part IV Carabids in human-managed habitats in a human-dominated era……...………………..67 1. Ground beetle assemblages in narrow hedgerows in a Danish agricultural landscape .............68 2. Ground beetle assemblages along an urbanisation gradient………………………..……...….77 3. The potential effect of transgenic plants on generalist natural enemies of pests: Tri-trophic interactions between two carabids and their prey fed on GM-product-containing diet………………………..……………………..………86 Discussion…………………………………………...………………………………………...…96 Part V – Theoretical studies……………………………………………...…..…………………103 1. The influence of matrix and edges on species richness patterns of ground beetles in habitat islands........................................................................................104 2. The collapse of animal assemblages in isolated forest fragments ...........................................108 3. The importance of life history in ecological theory - size trends in a ground beetle assemblage in relation to life history parameters....................................................111 4 Discussion....................................................................................................................................115 Part VI. Final thoughts: Ecology in an age of human domination...............................................121 Acknowledgements......................................................................................................................125 References cited...........................................................................................................................126 5 Introduction One of the famous sayings of modern entomology is provided by the evolutionary biologist, B.S. Haldane. At a debate with his theologist colleague, he was asked what he had discovered about the Lord during his lifetime of studying His creation. "That He has an inordinate fondness for beetles" replied Haldane with aplomb (Evans & Bellamy 1996). The basis of Haldane's statement is that the order of beetles, Coleoptera is the most spe- cies-rich on Earth. Within beetles, one can argue, the ground beetles, Carabidae must be one of His favourites, because among beetles, this family is one of the most speciose ones. It should be no surprise then, that the study of the family of ground beetles is also very popular. A search made on the Web of Science database resulted in >3000 publications, published between 1996 and 2005, that were dealing with ground beetles. This field of study therefore cannot be considered esoteric, the domain of a few special- ists. I started my research on carabids in 1978, one year after the premature death of the eminent German carabidologist, Hans-Ulrich Thiele, and the publication of his influential treatise on ground beetles (Thiele 1977) in which he summarised, in an almost encyclopaedic manner, our knowledge about the ecology of ground beetles. According to the then-current ecological para- digm, this book was mostly descriptive, with an emphasis on environmental physiology, culti- vated by Thiele himself at a high level. The following 30 years brought new ideas and rapid pro- gress, especially in ecology, and especially in Europe. Several researchers (e.g. Piet den Boer and his co-workers in the Netherlands) started their detailed, long-term studies that unearthed several important, so far unknown aspects of the ecology of ground beetles: the pattern and causes of long-term fluctuations of species, dispersal, population survival. These were laying the founda- tion for several new areas, including metapopulation theory, ecological restoration, and conserva- tion biology. Others extended the methodological toolkit, making it possible to study phenomena, even quantitatively, that were unavailable due to methodological limitations. These included the use of the harmonic radar in habitat use studies (Mascanzoni & Wallin 1986), the adoption of biochemical (isoenzyme activity, Lövei 1986) and immunological (ELISA, Crook & Sunderland 1984) methods to study feeding, the combination of laboratory and field methods in population biology (Baars & van Dijk 1984). My overall aim was to study the population and assemblage patterns of ground beetles living in human-influenced habitats. In our age, sometimes called "homogeocene", this meant more and more of our globe. Human influence is ever increasing, causing a homogenisation of the global flora and fauna ("Macdonaldisation", Lövei 1997), raising the global extinction rate by at least two orders of magnitude (Lövei 2007), and appropriating an ever-increasing share of globally available resources. These developments had a unifying influence on the study (also, but not only) of carabids. 6 This Thesis summarises my studies on ground beetles in five sub-areas. For any field of science, it is important to have periodical reviews, syntheses of known information. The first part is based on such a review (Lövei & Sunderland 1996), and briefly describes the main features of the ecology and behaviour of ground beetles. The second part presents a few methodological in- novations that may offer potential for the further development of the field. Among them there are field as well as data evaluation methods. The third part concerns life history studies. Knowledge of the biology of the study organ- isms in ecology is essential. Superficial knowledge in this regard can easily derail ecological studies (on such example is discussed in Lövei & Magura 2006), Therefore it is not only natural, but an obligation for an ecologists to contribute to our still-fragmentary knowledge on life histo- ries. This may be a near-endless task because we can hardly declare that we know "everything there is worth knowing" about a species. The current situation is, however, very differernt: a large part of the known arthropods are represented in collections by a single specimen (single- tons), about which we know nothing except the location of its single occurrence and species name (Allison & al. 1997.). With my colleagues, I have been involved in studying life history traits of ground beetles in Hungary, New Zealand an Denmark. Part III is based on this work. Part IV is devoted to studies on ground beetles, both at species and assemblage level, in habitats under anthropogenic influence. First results of studies with hedgerows are presented that show that forest faunal elements can be maintained in an agricultural landscape in Denmark. Sec- ond, the impact of an urbanisation gradient (forest- suburban forest – forest fragment in urban park) on ground beetles is explored. In order to analyse the impact of urbanisation on arthropod (ground beetle) diversity, species richness trends are not instructive – in our Danish study site, the most species rich site was the urban forest fragment. This, however, maksed several impor- tant effects of urbanisation on ground beetles. Included are studies linked to the environmental impact of genetically manipulated plants. The reason is that ground beetles are important in biological control, nutrient cycling, and linking above-and belowground ecological systems, and the genetically manipulated plants can influence these (positively or negatively). Consequently, it is important to include beneficial ar- thropods (including ground beetles) into environmental impact evaluations. Currently, there are few studies (of mostly short duration) concerning the impact of transgenic plants on biological control agents (Lövei & Arpaia 2005). The study of a single, well defined group is interesting in itself, but it is also important to link such results to theoretical knowledge. Results emerging from the study of ground beetles have been important to develop ecological theory (for example the long-term studies by den Boer (1987) that lead to the concept of metapopulations) albeit not to the extent that their abundance, wide occurrence and the amount of knowledge would justify. Results related to theoretical as- pects are summarised in the fifth part, which contains results
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