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IN a GRASSLAND HABITAT by DONALD

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IN a GRASSLAND HABITAT by DONALD THE ECOLOGY OF SOME SURFACE-DWELLING COLLEMBOLA (INSECTA) IN A GRASSLAND HABITAT by DONALD RUSSELL REYNOLDS, B. Sc. , A. R. C. S. A thesis submitted for the degree of Doctor of Philosophy in the Faculty of Science of the University of London MARCH 1974. Department of Zoology, & Applied Entomology, Imperial College Field Station, Silwood Park, Sunninghill, Ascot, Berks. 2 Abstract Some aspects of the biology of surface-dwelling Collembola are reviewed with emPhasis on adaptations to the surface mode of life. Possible methods of sampling for these animals are critically reviewed, and a new sampling method developed giving particular attention to the factors effecting'its efficiency. Population estimates have been made for four species of surface-dwelling Collembola over a period of two years, and for one of the species, Entomobrya nicoleti, a detailed analysis of the changes in the population size and structure in relation to various environmental factors was undertaken using regression methods. A little-used multivariate procedure (orthogonalized multiple regression) has been evaluated in this context. Tentative interpretations are made of the causal mechanisms underlying the statistical relationships revealed in the analyses. A laboratory and field investigation of the role of temperature on the development and survival of surface-dwelling Collembola, particularly the lethal effects of sub-zero temperatures, has been carried out. The dispersion patterns of the Collembola have been analysed by a variety of techniques, and the implications of the patterns for the ecology of the animals discussed. Finally, a methodological investigation of the applications of some clustering techniques to ecological data has been undertaken with emphasis on the behaviour of the test data under the various procedures. TABLE OF CONTENTS Page 1) Acknowledgements. 5 2) Introduction. 6 3) Taxonomic Note. 14. 4) Classification of Species Studied. 16 5) Review of Some Aspects of Biology: a) Habitat and Distribution. 17 b) Food. 25 c) Predation and Parasitism. 4.1 6) Description of the Study Area. 51 7) The Sampling of Surface-Dwelling Collembola: a) Description of Sampling Procedure. 54. b) Discussion of Methods of Sampling for Surface-Dwelling 57 Collembola. 8) Stage-Grouping. 84. 9) Collection. 86 10) Culturing. 87 11) Seasonal Abundance of the Four Species of Surface-Dwelling Collembola. 89 12) Analysis of Stage-Group Data: Life-Histories in the Field. 102 13) Regression Analysis: a) Methods. 115 b) Selection of Variables. 128 c) Analysis of Relationship between numbers of E. nicoleti, Suction-Sample Data, and Weather, by Orthogonalised Multiple Regression. 133 4 d) Analysis of Relationships between Winter populations of 14i. E. nicoleti and Weather. e) Analysis of Relationships between E. nicoleti populations And Weather: Main Analysis. 449 14) Egg Mortality in the Field. 156 15) The Analysis of Dispersion: a) Review. 163 b) Methods. 167 c) Results and Discussions: 1) Fitting of theoretical frequency distributions. 177 ii) Indices of Aggregation. 199 iii) Taylor's Power Law 2011. iv) Distribution Pattern as related to various Environmental Factors. 212 16) Cluster Analysis: a) Methods. 225 b) Results and Discussion of Single-Linkage Classifications. 231 c) Results and Discussion of Ordinations. 236 17) The Effects of Temperature: a) Relationship of Egg-Development Rate and Temperature. 259 b) Cold-Hardiness. 266 18) General Discussion. 291 19) Summary. 296 20) References. 299 ACIMOWLEDGEMENTS' This study has greatly benefit ed from stimulating discussions I have had with many colleagues in the Departments of Zoology and Botany at Imperial College, and also from their helpful advice and assistance on many occasions; to all who took an interest in the project, my thanks. In particular I am indebted to: Professor T. R. E. Southwood for the research facilities extended to me at Imperial College Field Station, Silwood Park; Mr. H. E. Goto for supervising the project, and for confirming the identifications of the Surface-dwelling Collembola; Mr. R. G. Davies for his invaluable assistance with various statistical aspects of the study; Drs. B. E. J. Wheeler and G. Jackson for advice on some mycological problems; Mr. A. Broodbank for his assistance in carrying out the sampling programme in the field. This study was made whilst I held a Natural Environment Research Council Studentship. Many of the analyses were carried out on the University of London's C. D. C. Computer using terminal facilities provided under a grant from the Ford Foundation. 6 INTRODUCTION The ecology of Collembola has been studied for over 60 years, since the early papers of Diem (1903) and Dahl (1912), and a large and scattered literature has accumulated. Salmon (1964) has compiled an extensive bibliography and many papers have been published on the subject since then. (See reviews by Paclt (1956), Christiansen (1964), Hale (in Burges and Raw 1967), Butcher, Snider and Snider 1971, and Healey (in Phillipson 1972)). However, many of the major studies in Continental Europe have been synecological or faunistic in nature, e.g. Gisin (1943, 1951), Janetschek (1949), Haarlov (1960), Nosek (1967), Cassagnau (1961), Cassagnau and Rouquet (1962),- Stebajeva (1963) and Rusek (1968). These biocoenological studies have been mainly concerned with distinguishing and classifying associations, or successions of associations and attempt- ing to "label" these by indicator species. This work is often combined with descriptions of various morphological—ecological features of the fauna which are adaptations to its special environment (life—form studies), see Macfadyen (1963 ch.13). Ecology in some countries is almost synonymous with work of this sort (Macdadyen 1963). These studies have given rise to a proliferation of definitions, and have produced descriptions of particular associations but they have, perhaps, been rather unsuccessful in analysing the nature of the dynamic effects of the various environ- mental agencies on the intrinsic attributes of the species which determine population size and quality at any given time and how these change with time. Peus (1954) (quoted by Clark et. al. 1967) makes the following criticisms of the biocenological approach; that it lacks acuity as an analytical method, that it is not convenient for experimental 7 investigations of numerical regulation and therefore it tends to preclude inductive synthesis. There has, of course, also been much work on the population ecology of Collembola and some justification may be required for embarking upon yet another study. Firstly, this investigation is only concerned with the surface-dwelling, arthropleonan species of the collembolan fauna. This group has been rather neglected compared with the soil-dwellers as Christiansen (1964) has pointed out. This may have been due partially to the difficulties in sampling the surface-dwelling forms as these species usually occur in small numbers in samples which were designed for the soil species. The study of surface-dwelling symphypleonan population dynamics has been greatly aided by the pest status of Sminthurus viridis and thus more is known about this group (Wallace 1967, 1968 and Walters 1968). While the present study was in progress the situation with regard to the surface-dwelling arthropleonans has been partly rectified by Joosse and her co-workers (1965 onwards). Secondly, many of the early studies were subject to some of the following drawbacks:- 1) Early workers were unavoidably impeded by the lack of efficient extraction methods, and until recently there has been little experimental work designed to elucidate the mechanisms behind the extraction methods (e.g. the behavioural responses of the animals under the various stimuli applied during extraction by dynamic-type methods). However, the recent papers of lief (1972, 1971), Vannier (1969, 1970, 1971), Macfadyen (1968) among others, lead one to suppose that there will soon be a logical basis for the development of more efficient extraction methods. 8 2)Another drawback was the limited period over which samples were taken in many of the earlier studies (1 year or even less). This meant that insufficient evidence was available as to whether the population curves presented were representative of regular annual cycles as was often implied by the authors. Dhillon and Gibson (1962) found that the populations sampled by them exhibited rather different trends in each of the years studied. The further analysis of populations to determine the main disturbing and regulating factors of course requires several years accurate census data (a minimum of five) for all stages of the animal, and none is available for a collembolan population (except that for Sminthurus viridis obtained by Wallace (1967). 3)Perhaps due to the synecological bias in collembolan ecology, and also to the fact that a single extraction method auto- , matically produces many species, there has been a tendency to investigate a large range of species rather than concentrate on a more detailed study of one or a few species combined with relevant experimental work. This has meant that the work effort has been spread rather thinly over the species involved. 4)A statistical approach has been lacking in many of the published studies. It is often difficult to know what level of precision was being aimed at in the sampling programme, and what con- fidence can be placed in the population estimates. It is also difficult to distinguish the significant fluctuations in population
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