Coleoptera: Cerambycidae) and Sapwood Moisture in a Drought Affected Northern Jarrah (Eucalyptus Marginata) Forest
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Behaviour of woodborers (Coleoptera: Cerambycidae) and sapwood moisture in a drought affected Northern Jarrah (Eucalyptus marginata) Forest Stephen A Seaton B.Sc. EnvSci, B.Sc. Biol Sci (Hons) Murdoch This thesis is presented for the degree of Doctor of Philosophy School of Veterinary and Life Sciences, Murdoch University October 2018 Declaration I declare that that to the best of my knowledge the work contained within this thesis is my own research, except where work by other sources are fully acknowledged, while I was enrolled in the degree of Doctor of Philosophy at Murdoch University, Western Australia. This thesis contains as its main content work which has not been previously submitted for a degree at any tertiary education institution. Stephen Seaton October 2018 ii Abstract A severe drought event during 2011 in the Northern Jarrah (Eucalyptus marginata Don Ex Smith) Forest of southwestern Australia, provided ideal conditions for an outbreak of the eucalyptus longhorned borer Phoracantha semipunctata Fabricius (Coleoptera: Cerambycidae). Located in a Mediterranean climatic zone, the jarrah forest region is expected to experience increased frequency of droughts causing tree mortality, yet little is known of endemic borer activity in response to tree drying. A detailed study was conducted within the jarrah forest following the drought, with the overall aim to provide an understanding of the role of sapwood moisture in controlling the infestation and development of longhorned borers. This was achieved using a field-based approach within a natural habitat and addresses the following key objectives: (a) assess the influence of seasons on infestation by key borer species, (b) determine the role of parasitoid wasps in controlling borer infestations, and (c) evaluate the influence of the two main Eucalyptus species, jarrah and marri (Corymbia calophylla L.A.S Johnson) on borer behaviour. Deterioration in crown health during the drought was associated with increased emergence of P. semipunctata and this was higher in larger lower tree sections where differences in emergence occurred between tree species and favoured marri with thicker bark compared to smaller higher tree sections with thinner bark. Poor survival of neonate larvae in both species was associated with overlapping galleries at high larval incidence, indicating intense competition for sapwood resources among developing larvae, while pupal survival remained high. This is the first study to show Coptocercus rubripes Boisduval to be an important borer along with P. semipunctata in the jarrah forest. Coptocercus rubripes mainly attacked in spring, prefers jarrah, has singular galleries per oviposition site, and grew slowly over spring to autumn, emerging in autumn–winter. Phoracantha semipunctata mainly attacked in the dry warm summer months, preferred marri, has many galleries per oviposition site, and grew rapidly over summer–autumn, emerging in summer the following year. Once sapwood moisture content in water stressed trees decreased by 19 % below turgid levels (independent of tree species), coinciding with a change in colour of canopy leaves of trees from green to yellow, conditions were suitable for borer larvae to establish. As sapwood dried, the larger P. semipunctata developed quicker and galleries were more extensive than iii C. rubripes. Larvae of various ages contributed to total sapwood consumption and reached a maximum at higher moisture contents in marri than jarrah. The native parasitic wasps Syngaster lepidus Brulle and Callibracon limbatus Brulle attacked maturing borer larvae of C. rubripes and P. semipunctata in stressed jarrah and marri hosts. Parasitism averaged 67 % and was inversely related to larval density. In the absence of parasitoids, borer survival in caged logs was greatly improved, although 30 % mortality still occurred and was attributed to increased competition among developing larvae. This research shows falling sapwood moisture has a key role for borer infestation and development in response to drought stress within forests and can be influenced by borer and tree species, parasitoids, and season. This knowledge will enable better management of cerambycids in forests subject to drought. iv Statement of Contribution of Others Chapter 2 of this thesis has been published in the journal ‘Forests’ titled Seaton, SA., Matusick, GB., Ruthrof, KB. & Hardy, GC. 2015. Outbreak of Phoracantha semipunctata in response to severe drought in a Mediterranean Eucalyptus forest. Forests, 6, 3868-3881. A was responsible for designing conducting and drafting and preparation of the manuscript for publication, B for identifying drought sites, assisting with fieldwork and manuscript editing, C principal supervisor for advice on design of experiments and critically revising the manuscript. Chapter 3-The contribution of George Matusick and Giles Hardy in design, field sampling, data analysis and editing of this chapter, is greatly appreciated. v Acknowledgements I greatly appreciate the continued enthusiasm, guidance, and suggestions on experimental work, statistical analysis and encouragement and support from my supervisors Professor Giles Hardy and Professor Bernard Dell (Murdoch University). I appreciate their expert knowledge and research experience, particularly in providing interaction and productive discussions. I would also like to thank Dr George Matusick and Dr Katinka Ruthrof (Murdoch University) for their mentoring, interactive discussions and support especially during the early stages of my thesis. The support of the Department of Biodiversity Conservation and Attractions (DBCA) for granting scientific permits in allowing access to the experimental forest sites. I would especially like to thank the Jarrahdale-shire manager for assisting with day to day management of sites during the project and Allan Wills for his encouragement and expertise in forest entomology during the project. Financial support given from the Holsworth Wildlife Research Endowment-Equity Trustees (2014–2017), an Australian Postgraduate Award and Murdoch University, made this work possible and is greatly appreciated. I greatly appreciate the assistance of Dr Bill Dunstan in field and laboratory work, and Ian McKiernan for the provision of glasshouse and oven facilities (both from Murdoch University). The support, camaraderie and encouragement of many colleagues and friends during my studies is greatly valued. Lastly, I would like to thank my family for their unwavering support, advice and encouragement during this journey. vi Table of Contents Declaration .......................................................................................................................................... ii Abstract ...............................................................................................................................................iii Acknowledgements ............................................................................................................................. vi Conference Presentations ................................................................................................................... xi Chapter 1 ............................................................................................................................................. 1 Introduction and literature review .................................................................................................... 1 Introduction ........................................................................................................................................ 1 Characteristics of the jarrah forest ................................................................................................... 1 Drought in the jarrah forest ............................................................................................................... 3 Soils ...................................................................................................................................................... 4 Drought, borers and occurrence of outbreaks .................................................................................. 5 Eucalyptus woodborers ....................................................................................................................... 6 Diversity ............................................................................................................................................... 6 Distinguishing features of Phoracantha semipunctata ..................................................................... 8 Life cycle .............................................................................................................................................. 9 Factors affecting borer behaviour ................................................................................................... 11 Sapwood moisture .............................................................................................................................. 11 Seasons ............................................................................................................................................... 12 Parasitoids .......................................................................................................................................... 12 Tree species and host tree environment ............................................................................................. 14 Conclusions .......................................................................................................................................