Community level consequences of adaptive management through Climate Matching: oak galls as a model system Frazer H. Sinclair Submitted for the degree of Doctor of Philosophy University of Edinburgh 2011 1 Declaration This thesis is submitted to the University of Edinburgh in accordance with the requirements for the degree of Doctor of Philosophy in the College of Science and Engineering. Aspects of the presented work were made possible by collaboration and data sharing with individuals and institutions, details of which are presented below. Chapter 2. The French National Institute for Agricultural Research (INRA) provided various phenotypic and genotypic data from oak provenance trials that are under their management. All presented analyses of these data are my own. Chapter 3. INRA allowed access to their established oak provenance trial at the forest of Petite Charnie in Sarthe, Northwest France. Insect surveys at the trial were conducted by me, and by volunteers under my supervision. All presented analyses of these data are my own. Chapter 4. Insect specimens were collected by me from the oak provenance trial at Petite Charnie with the permission of INRA. Approximately 1/3 of DNA extractions and PCR reactions were conducted by Konrad Lohse, Julja Ernst, and Juan Carlos Ruiz Guajardo. All presented analyses are my own. Chapter 5. Insect specimens were sourced from the Stone laboratory collections at the University of Edinburgh. Unpublished DNA sequence data from 6 parasitoid individuals were provided by Konrad Lohse. All presented analysis of this data is my own. Unless otherwise stated, the remaining work and content of this thesis are entirely my own. Signed: Frazer Sinclair 31/10/2011 2 Acknowledgments I am extremely grateful to my supervisors – Karsten Schonrögge, Graham Stone, and Stephen Cavers – for introducing me to the fascinating world of oaks and galls, and for their support throughout this project. I fear I may be hooked, and will still be dreaming of gallwasps long after the dust has settled on this thesis. This project benefited greatly from collaboration with the French National Institute for Agricultural Research (INRA), and so massive thanks must go to Alexis Ducousso, Rémy Petit, Antoine Kremer, Sophie Gerber, and Stefanie Wagner, for providing data from the Petite Charnie provenance trial and for their hospitality during various visits to Pierroton. At various stages of this project I have been aided by members of the international gall community, whose knowledge and enthusiasm for tiny wasps is inspiring. Many thanks therefore to George Melika, György Csoka, Dick Askew, Jose-Luis Nieves- Aldrey, Juli Pujade-Villar, Chang-Ti Tang, and Man-Maio Yang. Having started this PhD is blissful ignorance of phylogenetics, particular thanks must go to James Nicholls and Konrad Lohse for patiently teaching me from scratch. Thanks also to past and present Stone lab members for their advice and friendship along the way: Jack Hearn, Pablo Fuentes-Utrilla, Alex Hayward, Julja Ernst, Juan Carlos Ruiz Guajardo, Tracey Begg, Riikka Kaartinen, Laura Riggi, Marlous de Boer, Kasia Mikolajczak, and Sarah White. One million French galls require a lot of counting and I could not have done it without the help of the following fieldwork heroes: Kat Bradley, Julja Ernst, Matt Wainhouse, Phil Butterill, Stefan Curtis, Katherine Rushton, and even on occasion - Karsten Schonrögge. Thanks also to the fine folk of Bernay and Conlie – especially the Busson brothers, Philippe & Pierrette, and the Tabot family - who tolerated my shameful ignorance of the French language and made the time spent in France so enjoyable. Finally, my deepest gratitude to my parents, whose patience and support in the past year has been amazing, and to whom I dedicate this thesis. 3 Abstract In the present century, ecosystems across the globe will be subject to profound changes in climate. Forests are expected to be particularly sensitive to such change as the long life span of trees limits the potential for rapid adaptation. In order to preserve commercial viability and the essential ecosystem services provided by forests, there has been much interest in strategies for managing the adaptation of trees to their climatic environment. Climate Matching has emerged as one such strategy, whereby climate models are used to identify provenances – tree populations at a particular locality - with seed expected to be well adapted to the future conditions of a particular planting site. Debate continues about the feasibility and merit of this and other approaches, but it has yet to be demonstrated that the underlying assumptions of Climate Matching are valid for focal European tree species. Furthermore, a potentially major omission thus far has been consideration of how the Climate Matching strategy might influence associated organisms. Given the widely demonstrated bottom-up effects of foundation species genotype that have emerged from the field of community genetics, it is possible that planting seed of non-local provenance could effect forest organisms such as insect herbivores. In this thesis, I investigate the underlying assumptions of Climate Matching and its community level consequences using a model system of cynipid oak galls on Quercus petraea. Following a general introduction to Climate Matching and the study system, in Chapter 2 I use data from a provenance trial of Q. petraea in France to explore a central assumption of the Climate Matching strategy: that provenances of focal tree species show climate associated variation in adaptive phenotypic traits. In Chapter 3, I explore correlations between these phenotypic traits and the abundance, diversity, and community composition of an associated guild of specialist gall-inducing herbivores. Tree phenological traits in particular showed strong patterns of adaptation to climatic gradients, and influenced the abundance and community structure of galling species. However, as the response to non-local tree provenances was not strongly negative, it was considered unlikely that mixed planting of local and Climate Matched provenances would have sever impact on the gallwasp community. 4 Having assessed the bottom-up effects of provenance phenotypic variation on the galling community, my ultimate aim is to extend analysis to include associated hymenopteran inquilines and parasitoids. However, interpretation of effects at this level is hindered by taxonomic uncertainty, with a growing appreciation that morpho-taxa may not represent independently evolving lineages (i.e. ‘true’ species). In Chapters 4 & 5 I therefore develop approaches for addressing taxonomic uncertainty with this ultimate aim in mind. In Chapter 4, I apply a DNA barcoding approach to parasitoid and inquiline specimens reared from the provenance trial, and compare taxa based on barcodes with those based on morphology to identify points of taxonomic uncertainty. I also investigate the extent to which networks based on morphological and molecular taxa support contrasting conclusions of network properties. In Chapter 5 I explore the potential for molecular based resolution of species level taxonomic error in a challenging group of parasitoids: the genus Cecidostiba. Beginning with a framework of single locus DNA barcoding, I use data from multiple nuclear loci to reveal the existence of cryptic species. Finally, in Chapter 6 I explore the practicalities of Climate Matching in light of my empirical results, and suggest fruitful avenues for further research. 5 Contents Declaration…………………………………………………………………………...ii Acknowledgements………………………………………………………………….iii Abstract……………………………………………………………………………...iv Contents……………………………………………………………………………....1 Community level consequences of adaptive management through Climate Matching: oak galls as a model system ................................................................ 1 Declaration ............................................................................................................... 2 Acknowledgments ........................................................................................................ 3 Chapter 1 - Introduction ............................................................................................. 11 1.1. Forests and climate change in the UK............................................................. 11 1.1.1. Local adaptation to climate ...................................................................... 15 1.1.2. Community effects of tree provenance .................................................... 17 1.1.3. Provenance research ................................................................................. 18 1.2. Study system ................................................................................................... 20 1.2.1. Quercus petraea ....................................................................................... 20 1.2.2. Petite Charnie provenance trial ................................................................ 22 1.2.3. Oak gall communities .............................................................................. 23 1.3. Thesis outline .................................................................................................. 28 Chapter 2 - Are provenances of Quercus petraea locally adapted to climate? .......... 31 2.1. Introduction ..................................................................................................... 31 2.1.1. Local adaptation ......................................................................................