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Fungi and Their Potential As Biological Control Agents of Beech Bark Disease

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Fungi and Their Potential As Biological Control Agents of Beech Bark Disease Fungi and their potential as biological control agents of Beech Bark Disease By Sarah Elizabeth Thomas A thesis submitted for the degree of Doctor of Philosophy School of Biological Sciences Royal Holloway, University of London 2014 1 DECLARATION OF AUTHORSHIP I, Sarah Elizabeth Thomas, hereby declare that this thesis and the work presented in it is entirely my own. Where I have consulted the work of others, this is always clearly stated. Signed: _____________ Date: 4th May 2014 2 ABSTRACT Beech bark disease (BBD) is an invasive insect and pathogen disease complex that is currently devastating American beech (Fagus grandifolia) in North America. The disease complex consists of the sap-sucking scale insect, Cryptococcus fagisuga and sequential attack by Neonectria fungi (principally Neonectria faginata). The scale insect is not native to North America and is thought to have been introduced there on seedlings of F. sylvatica from Europe. Conventional control strategies are of limited efficacy in forestry systems and removal of heavily infested trees is the only successful method to reduce the spread of the disease. However, an alternative strategy could be the use of biological control, using fungi. Fungal endophytes and/or entomopathogenic fungi (EPF) could have potential for both the insect and fungal components of this highly invasive disease. Over 600 endophytes were isolated from healthy stems of F. sylvatica and 13 EPF were isolated from C. fagisuga cadavers in its centre of origin. A selection of these isolates was screened in vitro for their suitability as biological control agents. Two Beauveria and two Lecanicillium isolates were assessed for their suitability as biological control agents for C. fagisuga and nine Trichoderma isolates were screened for their antagonistic ability against Neonectria spp. Colonisation of beech saplings with Beauveria, Lecanicillium and Trichoderma isolates was attempted using three inoculation techniques. 3 TABLE OF CONTENTS ABSTRACT ...................................................................................................................... 3 TABLE OF CONTENTS .................................................................................................. 4 LIST OF FIGURES .......................................................................................................... 7 LIST OF TABLES .......................................................................................................... 10 LIST OF APPENDICES ................................................................................................. 11 ACKNOWLEDGEMENTS ............................................................................................ 12 GLOSSARY .................................................................................................................... 13 1. GENERAL INTRODUCTION ............................................................................... 14 1.1 INVASIVE SPECIES ........................................................................................... 14 1.2 THE GENUS FAGUS (FAGACEAE) BEECH .................................................... 15 1.2.1 Fagus sylvatica (L.), European beech ............................................................ 15 1.2.2 F. grandifolia (Ehrhart), American beech ...................................................... 18 1.3 PESTS AND DISEASES OF FAGUS SPP. ......................................................... 18 1.4 BEECH BARK DISEASE (BBD) ........................................................................ 19 1.4.1 Cryptococcus fagisuga (Lindinger), Beech scale .......................................... 20 1.4.2 Neonectria species ......................................................................................... 21 1.4.3 Abiotic factors affecting the development of BBD........................................ 28 1.4.4 Incidence and distribution of BBD ................................................................ 28 1.4.5 Impact of BBD on forest composition ........................................................... 29 1.5 CONTROL METHODS ....................................................................................... 31 1.5.1 Biological control ........................................................................................... 31 1.6 FUNGAL ENDOPHYTES ................................................................................... 34 1.7 MYCOBIOTA OF FAGUS SYLVATICA .............................................................. 36 1.8 ENTOMOPATHOGENIC FUNGI (EPF) ............................................................ 37 1.9 OUTLINE AND OBJECTIVES OF THE PHD THESIS ..................................... 38 2. SURVEYS FOR FUNGAL STEM ENDOPHYTES OF EUROPEAN BEECH (FAGUS SYLVATICA) IN THE UK AND ARMENIA AS POTENTIAL CLASSICAL BIOLOGICAL CONTROL AGENTS OF BEECH BARK DISEASE .......................... 40 2.1 INTRODUCTION ................................................................................................ 40 2.1.1 Fungal endophytes ......................................................................................... 40 2.2 MATERIALS AND METHODS .......................................................................... 44 2.2.1 Survey for endophytes in the UK ................................................................... 44 2.2.2 Survey for endophytes in Armenia ................................................................ 47 2.2.3 Isolation, storage and identification of endophytes ........................................ 47 2.2.4 Data analysis of UK endophytes .................................................................... 49 2.2.5 Molecular identification of selected endophytes............................................ 50 2.3 RESULTS ............................................................................................................. 51 4 2.3.1 Total endophyte isolates ................................................................................. 51 2.3.2 UK endophytes ............................................................................................... 57 2.3.3 Armenian endophytes..................................................................................... 60 2.3.4 Molecular identification of selected endophytes............................................ 60 2.4 DISCUSSION ....................................................................................................... 62 3. ENTOMOPATHOGENIC FUNGI ASSOCIATED WITH BEECH SCALE (CRYPTOCOCCUS FAGISUGA) IN ARMENIA AND A PRELIMINARY ASSESSMENT OF THEIR POTENIAL FOR BIOLOGICAL CONTROL .................. 66 3.1 INTRODUCTION ................................................................................................ 66 3.1.1 Entomopathogenic fungi ................................................................................ 66 3.2 MATERIALS AND METHODS .......................................................................... 69 3.2.1 Survey of entomopathogenic fungi (EPF) in Armenia .................................. 69 3.2.2 Entomopathogen collection and isolation ...................................................... 69 3.2.3 Morphological identification of EPF ............................................................. 69 3.2.4 Development of a C. fagisuga inoculation technique .................................... 70 3.2.5 Molecular analysis of endophytic Beauveria isolates .................................... 71 3.2.6 Radial growth of Lecanicillium, Beauveria and Chaunopycnis isolates........ 75 3.2.7 Data analysis .................................................................................................. 76 3.3 RESULTS ............................................................................................................. 77 3.3.1 Collection and culture of EPF in Armenia ..................................................... 77 3.3.2 Inoculation of C. fagisuga eggs ..................................................................... 80 3.3.3 Inoculation of C. fagisuga juveniles .............................................................. 82 3.3.4 Molecular analysis of endophytic EPF .......................................................... 84 3.3.5 Radial growth of Beauveria, Chaunopycnis and Lecanicillium isolates........ 88 3.4 DISCUSSION ....................................................................................................... 91 4. IN VITRO SCREENING OF ENDOPHYTIC TRICHODERMA ISOLATES FOR FUNGAL ANTAGONISM OF NEONECTRIA COCCINEA AND N. FAGINATA ...... 95 4.1 INTRODUCTION ................................................................................................ 95 4.1.1 Morphology and taxonomy of the genus Trichoderma ................................. 95 4.1.2 Ecology of Trichoderma spp. ......................................................................... 96 4.1.3 Trichoderma spp. as biological control agents .............................................. 97 4.1.4 Mechanisms of fungal antagonism................................................................. 98 4.2 MATERIALS AND METHODS ........................................................................ 101 4.2.1 Fungal cultures and their maintenance ......................................................... 101 4.2.2 Growth rates of Neonectria isolates ............................................................. 102 4.2.3 Measurement of competition through dual plate interactions between Neonectria and Trichoderma isolates ..................................................................
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