595.762.12:591.5:575.8 MISCELLANEOUS PAPERS 18 (1979) AGRICULTURAL UNIVERSITY WAGENINGEN THE NETHERLANDS ON THE EVOLUTION OF BEHAVIOUR IN CARABID BEETLES Report of a symposium, held at the fieldstation Rees-Grietherbusch of the Zoological Institute of the University of Cologne September 10-13, 1978 edited by P. J. DEN BOER, H. U. THIELE and F. WEBER H. VEENMAN & ZONEN B.V. - WAGENINGEN - 1979 11 r cpt,j O 1 We dedicate these papers to the memory of Carl H. Lindroth (8.9.1905 - 23.2.1979), who inspired us all CONTENTS PREFACE 1 INSTEAD OF AN INTRODUCTION 3 PARTICIPANTS 4 1. Behaviour and body structure P. Nagel : The classification of Carabidae 7 P. Nagel : Aspects of the evolution of myrmecophilous adaptations in Paus- sinae (Coleoptera, Carabidae) 15 P. Brandmayr & T. Zetto-Brandmayr : The evolution of parental care phenomena in Pterostichine ground beetles with special reference to the genera Abax and Molops (Coleoptera, Carabidae) 35 2. Behaviour and ecophysiological functions H. U. Thiele: Intraspecific differences in photoperiodism and measurement of day length in Pterostichus nigrita PAYKULL (Coleoptera, Carabidae) 53 G. Leyk : Intraspecific differences in the activity rhythms of the carabid beetle Pterostichus nigrita PAYKULL 63 G. Lamprecht & F. Weber : The regressive evolution of the circadian system controlling locomotion in cavernicolous animals 69 C. Juberthie : L'évolution des coléoptères Trechinae souterrains (Coleoptera, Carabidae) 83 3. Behaviour and life-history phenomena D. Mossakowski: Evolution of habitat preference illustrated by the phylo- geny of Chrysocarabus (Coleoptera, Carabidae) 103 W. Paarmann : A reduced number of larval instars as an adaptation of the desert carabid beetle Thermophilum (Anthia) sexmaculatum F. (Coleo­ ptera, Carabidae) to its arid environment 113 W. Paarmann : Ideas about the evolution of the various annual reproduction rhythms in carabid beetles of the different climatic zones 119 T. Bauer: The behavioural strategy used by imago and larva of Notiophilus biguttatus F. (Coleoptera, Carabidae) in hunting Collembola 133 4. Behaviour and the dynamics of populations P. J. den Boer: Populations of carabid beetles and individual behaviour: General aspects 145 P. J. den Boer : The individual behaviour and population dynamics of some carabid beetles of forests 151 Th. S. van Dijk: Reproduction of young and old females in two carabid beetles and the relationship between the number of eggs in the ovaries and the number of eggs laid 167 P. J. M. Mols: Motivation and walking behaviour of the carabid beetle Pterostichus coerulescens L. at different densities and distributions of the prey, a preliminary report 185 T. H. P. van Huizen : Individual and environmental factors determining flight in carabid beetles 199 Some remarks in retrospect (P. J. den Boer) 213 PREFACE At a symposium like this one, in most cases people come together who are engaged in the same field of science, quite independent from the animal species or group from which they derive their insights. In extreme cases, this spectrum may reach from protozoa to mammals. It is also attractive, however, if scientists meet who, using different methods and having different starting points, are occupied with the same group of animals, as we are with carabids. Of course, a unifying concept and a common interest have to bring us together. It seems that all of us who have gathered here are united by an interest in the question as to how the great variety of species of carabids has come into existence. If we are convinced that the theory of evolution is the unifying concept of biology par excellence, one feels that it is a most exciting task to trace the evolution of carabids. With something like 40,000 species, they are really a phylogenetic 'success', perhaps the greatest after the Curculionidae. It seems that the variation in carabid behaviour surmounts that of their external properties. It is not especially difficult to teach a child what a carabid is. You need only exclude a few cases. But the variety in behaviour of the carabids is very great. Those differences in behaviour are the theme of this symposium. Discussing these topics, we must bear in mind the question of whether we are able, with our investigations, to contribute to the better understanding of the evolution and the history of geographical distribution in a great and important group of animals. Most of the scientists who are concerned with the biology of carabids started investigating the problem of habitat binding or choice of habitat of our beetles. This applies also to most of the participants of this symposium. What binds carabid species to their often narrowly restricted habitats? From the early work of CARL LINDROTH, the founder of experimental ecology of carabid beetles, it became obvious that, in addition to the physiological capability, behaviour is decisive for the adaptation of a species to its special mode of life. Nowadays, we know that besides a preference for special microclimatic conditions, the following complexes of behaviour guarantee the annidation of species : brood care, food choice, mobility, diurnal rhythm of activity, and annual periodicity of development. All of these patterns of behaviour are genet­ ically variable. They were first investigated as if they were stable and unvariable characteristics of the species. But today more and more we investigate the question of how they became altered in the course of the genetic adaptation of populations, the formation of subspecies and species, and in phylogeny. It is the goal of this symposium to gain some insight into these processes. We are convinced that the value of the individual investigations is increased by a combined publication under a comprehensive heading. It has already become a good tradition that the Landbouwhogeschool Wageningen, The Netherlands, opens its 1 excellently equiped journal 'Miscellaneous Papers' to voluminous papers on carabid ecology. As early as 1971, the Symposium 'Dispersal and Dispersal Power of Carabid Beetles', held at the Biological Station in Wijster, was published, as well as den Boer's work on 'Dispersal Power and Survival' in 1977. The publication of this volume was made possible by numerous subscriptions from the Deutsche Forschungsgemeinschaft and the participants of this symposium. We are greatly indebted to them all. H. U. THIELE F. WEBER 2 INSTEAD OF AN INTRODUCTION The double role played by modes of behaviour in animal evolution has been pointed out by MAYR (1970). 1. Inborn behaviour is just as much exposed to the influence of evolutionary factors as are physical characteristics and undergoes the same kind of changes in the course of evolution. WICKLER (1970) regards behaviour 'so to say, as the most malleable organ of the organism ... The 'plasticity' of behaviour makes it especially useful as an adaptive feature and is one reason why behaviour often acts as 'pacemaker' in evolution'. This implies 'that, in the process of adaptive evolution, alterations in behaviour precede changes in body structure...'. 2. Considerable emphasis is placed by MAYR (1970) on the role of individual modifi­ cations in behaviour as causal factors in evolution. He writes: 'The particular signifi­ cance of behaviour as an evolutionary factor lies in the possibility offered to the individual of escaping certain environmental influences and of confronting others as they arise'. We mentioned above that this holds true to a particularly high degree for carabids. Thanks to their powers of activity they are readily able to move on to new habitats that present them with changed conditions of selection, initiating new evo­ lutionary processes and the occupation of new ecological niches. In this connection Mayr considers it to be a 'somewhat unexpected' discovery that 'the aspects of behaviour that have been especially thoroughly studied by ethologists, i.e. inter- and intraspecific relationships... are just those that play a relatively in­ significant role in macroevolution... Intraspecific behaviour consists mainly in an exchange of signals, and serves the purpose of communication or of misleading an enemy, or assists escape. Only rarely does it play a role in the discovery of a new ecological niche or adaptive zone (present author's italics). There is good reason for the ethologist's reluctance to concern himself with the ecological aspects of behaviour : they are extremely difficult to analyse. On the one hand, they often vary considerably even within one and the same population, and on the other, they always involve a strong, non-hereditary component. This is true of choice of food, choice of biotope and also of certain types of movement. The significance of types of behaviour in the larger evolutionary events becomes obvious when one considers the transition from aquatic to terrestrial life, from tree-life to one on the ground, or from an earthbound life to one involving flight'. The adaptive role of these types of behaviour in cladogenesis and speciation of any one animal group has seldom been the subject of such extensive investigation as in the carabids, and the studies on this group provide the zoologist with the means of bridging the gaps between ecology, behaviour and evolution. (This text is derived from THIELE, Carabid Beetles in Their Environments, 1977, p. 326-327) 3 PARTICIPANTS J. ALTHOFF: Zoologisches Institut der Universität Münster, Germany Ir. F. BALK: Biologisch Station Wijster, The Netherlands Dr. TH. BAUER: Zoologisches Institut der Universität Ulm, Germany Dr. P. J. DEN BOER: Biologisch Station Wijster, The Netherlands Dr. P. BRANDMAYR: Istituto di Zoologia e Anatomia Comparata, Università di Trieste, Italy Dr. TH. S. VAN DIJK: Biologisch Station Wijster, The Netherlands L. ERBELING: Zoologisches Institut der Universität Münster, Germany M. HEINE: Zoologisches Institut der Universität Münster, Germany Ing. T. H. P. van Huizen: Biologisch Station Wijster, The Netherlands. Dr. G. JUBERTHIE : Laboratoire Souterrain, CNRS, Moulis, France Dipl. Biol. D. KOCH: Zoologisches Institut der Universität Köln, Germany G. LEYK: Zoologisches Institut der Universität Köln, Germany W.