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By Andrew M. Brewer Thesis Submitted to the University Of INTERACTIONS BETWEEN DEMOGRAPHIC RATES, POPULATION DENSITY AND THE ENVIRONMENT - THE SPATIAL STRUCTURE OF THE RANGE OF THE HOLLY LEAF-MINER by Andrew M. Brewer Thesis submitted to the University of Sheffield for the degree of Doctor of Philosophy in the Department of Animal and Plant Sciences. NOVEMBER 2000 Summary Generalisations about the abundance structure of species' geographic ranges may have implications within a number of areas of applied ecology. However, empirical data is largely restricted to a single taxonomic group within one geographic region. One approach to the study of range structure and its ecological consequences is to examine the relationship between demographic rates and environmental conditions. However, most studies of population processes are at small spatial scales and it is not known to what extent patterns found at these scales may be extrapolated. This thesis addresses these issues using surveying techniques to measure spatial structure in both the densities and demographic rates of the holly leaf-miner (Phytomyza ilicis Curt.) at a wide variety of spatial scales. Geostatistical tools are used to analyse the data collected. At a regional scale, spatial structure in leaf-miner densities is apparent despite considerable variation between hosts within individual habitat patches. This structure can in part be accounted for by variation in habitat and altitude. Over the entire range, broad scale trends in population density can be detected which can also, be correlated with environmental variation.' Many demographic rates components exhibit spatial structure. However, their relationships both with population density and environmental variation are more complex. Per capita mortality rates did not correlate well with population density at any spatial scale. The difficulties inherent in relating population density with demographic rates and regulation by natural enemies are discussed. The demographic approach has also been used to explain.the positive interspecific abundance-distribution relationship. Computer simulation techniques are employed to explore this model. The results indicate that the positive relationship should be extremely robust under a high degree of variation between the demographic rates. However, data from the holly leaf-miner surveys suggest that current. models of range structure make unrealistic assumptio~s about environmental variation. The key to understanding range structure and its consequences may lie in our ability to make. generalisations about environmental structure. 11 Acknowledgements This project was financially supported by the Alfred Denny Studentship at the University of Sheffield. The European holly leaf-miner survey was funded by the Natural Environment Research Council. I am indebted to a number of people for their assistance during this project. Thanks to Melody McGeoch for help in the fieldwork and for useful discussion during the Sheffield Surveys. Many thanks to Rob and Kerry Briers, Sian Gaston, Trond Hofsvang, Jens Rolff, Phil Warren and most especially John Maryal for help and advice during the European survey. Thanks also to Bob Holt for an extremely helpful day of discussion about the vital rates model. I am particularly grateful for the support of my supervisor Kevin Gaston, who has been a continual source of inspiration throughout the project. This thesis is dedicated to my parents. 111 Declaration Two chapters in this thesis were jointly authored. The contributions I have made to these chapters are detailed below.. Chapter 4: Brewer A.M. & Gaston K.J. (manuscript) The population density structure of the geographic range of the holly leaf-miner. Chapter 5: Brewer A.M. & Gaston K.J. (manuscript) Spatial structure in the demographic rates of the holly leaf-miner across its geographic range. These manuscripts have been prepared for submission to Gikas and are based on an original idea by KJG. Data. throughout Europe were collected primarily by AMB with the help of a research assistant. AMB also created the database, conducted all data analyses and drafted the manuscripts. lV Table of Contents Chapter 1 Introduction.......................................................................................... 1 Chapter 2 The holly leaf-miner system ................................................................. 15 Chapter 3 The spatial structure of holly leaf-miner populations and variation in mortality rates at a regional scale....... .......................................... ...... .......... ........ 28 The structure of the geographic range of the holly leaf-miner.............................. 56 Chapter 4 The population density structure of the geographic range of the holly leaf-miner .............................................................................................................. 57 Chapter 5 Spatial structure in the demographic rates of the holly leaf-miner across its geographic range .................................................................................... 78 Chapter 6 The interspecific abundance-distribution relationship and the vital rates model ............................................................................................................ 104 Chapter 7 Discussion ......................................................................... ;.................. 143 References .................................................................................... ......................... 156 Appendices Program source code and documentation..... ............. ......... ....... ....... 169 Appendix 1 Spatial autocorrelation analysis program ....................... :................... 170 Appendix 2 Vital rates model simulation ............................................................. 184 v Chapter 1 Introduction HAs we cannot know the ,reason for a fact before we know the fact. We cannot know what a thing is before knowing that it is. " Aristotle The aims of population ecology are to describe patterns in the distribution and abundance of organisms through space and time, and to explain these patterns in terms of their underlying mechanisms. Much of our understanding of population processes is based upon research conducted at small spatial scales (Brown et al. 1995), typically at the level of individual populations confined to a single habitat patch. It has not been established whether the information collected from these studies can be applied to larger spatial scales. This is an important issue, since many of the environmental problems we face today are related to processes operating at these broad scales. The effects of global climate change, shifts in land-use and biological invasions on the current distribution and abundance of species is likely to have important consequences for agriculture, the conservation of biodiversity and the spread of disease. Macroecology (Brown & Maurer, 1989) is a relatively recent attempt to understand large scale ecological phenomena using the substantial database that has already been assembled on many characteristics of plant and animal populations. Where traditional ecological studies have largely focused upon the roles of individual organisms in community and population processes, macroecology aims to identify and explain patterns that become apparent when comparing species characteristics within assemblages. Typical units of study are the size and shape of the geographic range, mean population density and the mean body size of species. The approach has met with some success. A number of statistical relationships between such characteristics have been identified and some historically well-known patterns have received renewed attention and attempts at explanation (e.g. Gaston, 1994; Brown, 1995). However, it can be difficult to distinguish between competing hypotheses. Experimental work is seldom practical at these taxonomic and spatial scales and collection of the additional data required may be difficult and expensive. 1 For example, there has recently been a renewed attempt to make statistical generalisations about the internal abundance structure of the geographic ranges of species. If such generalisations could be made there would be obvious consequences within several areas of applied ecology such as the identification of high risk areas for pest outbreaks and the design and location of nature reserves. A number of intuitively­ appealing assertions have been made about range structure and its causes (e.g. Brown, 1984; Hengeveld, 1989; Maurer & Brown, 1989) but these are derived from just a few empirical studies which have been based on narrow taxonomic and geographic domains. One potentially useful approach to the investigation of range structure is given by Maurer & Brown (1989). They propose that it may best be understood in terms of changes across space in the four demographic rates that underlie population regulation; births, deaths, immigration and emigration. Birth and immigration sum to a net rate of population gain and death and emigration sum to a net rate of population loss. Where population gain and population loss intersect there is a stable equilibrium point. The general spatial pattern of population abundance may therefore be characterised as the set of equilibrium points of local popUlations across space. If the demographic rates are dependent upon local environnlental conditions, then range structure should ultimately reflect the spatial structure of the envirorunent. Maurer & Brown (1989) make a number of assumptions about the nature of envirorunental structure
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