Biology and Conservation Ecology of Selected Saproxylic Beetle Species in Tasmania‘S Southern Forests
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Biology and conservation ecology of selected saproxylic beetle species in Tasmania‘s southern forests Belinda Yaxley B.Sc. (Hons.) School of Zoology, University of Tasmania A research thesis submitted in fulfilment of the requirements for the degree of Doctor of Philosophy October 2013 Declaration I hereby declare that this thesis contains no material which has been accepted for the award of any other degree or diploma in any University, and to the best of my knowledge contains no copy or paraphrase of material previously published or written by any other person, except where due reference is made in the text of this thesis. Belinda Yaxley University of Tasmania 1st October 2013 Authority of Access This thesis is available for loan via the University of Tasmania and Forestry Tasmania. Copying any part of this thesis is prohibited for one year from the date this statement is signed; thereafter, copying is permitted in accordance with the Copyright Act 1968. Belinda Yaxley 1st October 2013 Chapter 4 is presented as a manuscript and was submitted for publication to Biodiversity and Conservation on the 30th of September 2013. The three authors and their subsequent contribution to the manuscript (shown as a percentage) are as follows: Belinda Yaxley – University of Tasmania, School of Zoology (60%) Initiation Experimental design Collection of samples Preparation of samples Manuscript writing Morag Glen – University of Tasmania, Tasmanian Institute of Agriculture (30%) Preparation of samples Analysis of samples Manuscript writing Simon Grove - Tasmanian Museum and Art Gallery (10%) Initiation Manuscript writing 2 ACKNOWLEDGEMENTS First and foremost I would like to thank my supervisors Dr. Simon Grove, Dr. Peter McQuillan, Assoc. Prof. Alastair Richardson and Dr Simon Wotherspoon for whose expertise and guidance I am ever grateful. I would also like to thank my family and friends who provided continual support and encouragement throughout the thesis process. A special thank you to volunteers who provided assistance in the field and to the staff at Forestry Tasmania. I greatly appreciate the assistance of specialists Dr. Morag Glen (University of Tasmania), Dr. Tim Wardlaw (Forestry Tasmania), Dick Bashford (Forestry Tasmania), Dr. Stephen Harris (Department of Primary Industries Parks Water and Environment), Chris Spencer, Dr. Karen Richards (Department of Primary Industries, Parks, Water and Environment) and Dick Chuter. I dedicate this thesis in loving memory of my grandfather James W. G. Yaxley. 3 ABSTRACT Many saproxylic beetle species (those associated with dead wood) are considered to be threatened by intensive forest management, yet few have been subjected to autecological studies that might help in their conservation. For this thesis, two lines of research were followed to investigate the autecology of six species thought to be vulnerable to intensive forestry. The study was carried out in the wet-eucalypt production forests of southern Tasmania. The first line of research explored methodologies that could be used to help understand life-history and feeding traits of log-dwelling beetles, in order that the resultant autecological information might be used as input into models of species persistence. Study 1a investigated the sexual development and life-history strategies of two sympatric species of stag-beetle, Lissotes menalcas and L. cancroides; and study 1b used molecular approaches to explore the feeding relationship of a mycophagous saproxylic beetle Prostomis atkinsoni and its putative food source, ‗gingerbread rot‘. Both studies in this line of research provided useful methodological insights. Additionally, the findings from study 1asuggest that coexistence of L. menalcas and L. cancroides may be enabled by their differing patterns of sexual development across seasons; while the findings of study 1b were equivocal, in that numerous fungal species were detected in the gut-contents of adult and larval P. atkinsoni but few were found in both; furthermore, there was little overlap in the mycota of beetles' guts and gingerbread rot. Unexpectedly, some species of basidiomycete known to be wood-rotters were found in the guts but not in the gingerbread rot itself. The second line of research comprised predictive modelling approaches at the level of (a) microhabitat (coarse woody debris) and (b) site and landscape, both aimed at identifying the habitat requirements of all six species and exploring the possibility of making reasonable predictions about the species‘ occurrence at each of these spatial scales. In study 2a, models performed well in relating the probability of occurrence of each of the six beetle species to the presence or absence of particular suites of rotten-wood types. The addition of site-scale variables did not greatly enhance the models‘ abilities to predict occurrence of the six species of beetles, but at this level, forest stand structure and age of 4 regeneration generally contributed the most explanatory power. In study 2b, geology, aspect, and time since last fire were good predictors of beetle occurrence across the study-area, although predictive ability may have been inflated by overfitting and autocorrelation. Overall, the models indicate that in unharvested forest these beetles tend to preferentially occur (within their chosen rotten-wood types) in multi-aged stands with a mature eucalypt element, while their continued occurrence in forest regenerating following clearfelling may be explained by their occupation of ‗legacy‘ coarse woody debris derived from the previous unharvested stand. The findings from both lines of research provide clear pathways for future exploration and modelling of species persistence in production-forest landscapes, and could help guide management practices accordingly, particularly in relation to the required spatial and temporal arrangement of mature forest. 5 ACKNOWLEDGEMENTS .................................................................................. 3 ABSTRACT ........................................................................................................... 4 CHAPTER 1 GENERAL INTRODUCTION ..................................................... 1 1.1.1 What is dead wood? ............................................................................................................... 2 1.1.2 Why is dead wood so important? ......................................................................................... 2 1.1.2.1 Dead wood as a carbon store ........................................................................................... 2 1.1.2.2 Dead wood as a slow-release fertiliser ............................................................................. 3 1.1.2.3 Dead wood as habitat for wildlife .................................................................................... 4 1.1.3 Different species of saproxylic beetles require different microhabitats in dead wood .... 5 1.2. EUROPEAN PERSPECTIVES ON THE ECOLOGY AND MANAGEMENT OF DEAD WOOD AND THE CONSERVATION OF SAPROXYLIC SPECIES. .................................................................................... 7 1.2.1 Why is conservation and management of dead wood worthy of focussed study? ............ 7 1.2.1.1 Current abundance of dead wood is no guarantee of future abundance ........................... 7 1.2.1.2 Loss of dead wood means a decline in saproxylic species ............................................... 8 1.2.1.3 Changes in dead-wood management ................................................................................ 9 1.2.1.4 Are current management recommendations for dead wood effective? .......................... 10 1.2.1.5 Dead wood in Australia ................................................................................................. 11 1.3 MANAGING DEAD WOOD IN TASMANIA: A CASE STUDY ...... 12 1.3.1 Same dead-wood thinking, different dead-wood situation ......................................... 14 1.3.2 Wet-eucalypt forest dynamics........................................................................................ 15 1.3.3 Forestry practices in Tasmania’s wet-eucalypt forests ............................................... 17 1.3.4 Biogeography and dead-wood habitat ......................................................................... 18 1.3.5 Thesis structure and objectives ...................................................................................... 19 1.3.6 Detailed overview of data chapters and synthesis ........................................................ 20 1.4 References........................................................................................................................ 24 CHAPTER 2 DETAILS OF STUDY AREA AND SPECIES ......................... 35 2.1 Description of study area and its forest types ...................................................................... 35 2.1.1 Location ............................................................................................................................ 35 2.1.2 Environmental features of the study area .......................................................................... 36 2.1.3 Management history of the study area .............................................................................. 37 2.1.4 Study Sites ........................................................................................................................ 39 2.2 Study species ..........................................................................................................................