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Conservation of Australian Insectivorous Marsupials

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Conservation of Australian insectivorous marsupials: biogeography, macroecology and life history Rachael A. Collett BSc Hons A thesis submitted for the degree of Doctor of Philosophy at The University of Queensland in 2018 School of Biological Sciences ABSTRACT The insectivorous marsupial Family Dasyuridae is endemic to Australia and Papua New Guinea. Dasyurids have a variety of life history strategies and are found in a wide range of habitats, latitudes and elevations. This makes them useful for exploring life history evolution, biogeographical patterns and macroecological theory. The life history and persistence of dasyurids are thought to be tightly linked to the distribution of their arthropod prey in space and time, but we currently have a limited understanding of the mechanisms. Arthropods are declining globally, and evidence suggests that climate change is already linked to changes in arthropod availability in the northern hemisphere. Understanding the relationship between insectivorous marsupials and their arthropod prey has become particularly important, as in 2018 two species from the dasyurid genus Antechinus were federally listed as endangered, and populations of other dasyurids appear to be declining across Australia. The aim of this thesis was to combine biogeography, macroecology and life history to determine what is influencing the distribution and persistence of Australian insectivorous marsupials from the Family Dasyuridae, with a focus on antechinus species in Queensland. I addressed evolution, rarity patterns and phenological relationships between dasyurid marsupials and their arthropod prey. Chapter 2 investigated how life history traits co-vary and are related to climatic variables, specifically how the evolution of dasyurid life history is related to productivity and predictability, and how environmental drivers are linked to patterns of life history variability. Based on analysis of data on life history characteristics of female dasyurids throughout Australia, there was evidence of two life history axes. The reproductive output axis including litter size and supernumerary young was related to annual rainfall and productivity, while position on the semelparity – iteroparity axis was associated with predictability of rainfall and arthropod availability. These results suggested how climate change might affect dasyurid marsupials and provide a broader understanding of their life history evolution that can be applied in a global context. Previous global and Australian studies have found that population density in mammals is lower in the tropics than at temperate latitudes. Chapter 3 tested whether dasyurids show a latitudinal gradient in population density, focusing on antechinus population density across latitudes and elevations in Queensland. Data from the literature, combined with intensive trapping and arthropod sampling showed that dasyurids were rarest in the tropics i and that antechinus population density was lowest at low latitudes and elevations. Population density was constrained by winter arthropod availability, which was associated with winter rainfall. High elevation sites maintained high arthropod availability throughout the year because they received orographic rainfall. These results help to explain rarity of antechinus species in the seasonal subtropics and tropics, including the Endangered silver-headed antechinus (A. argentus) in central Queensland, and the restricted Atherton antechinus (A. godmani) and rusty antechinus (A. adustus) in far north Queensland. The results also highlight the importance of Australian mountain-tops as refuge areas for insectivorous marsupials. Chapter 4 focused on timing of breeding and distribution of antechinus species across mountains in Queensland. I used the same data collection approaches as in chapter 3 to determine timing of breeding, evolutionary age of antechinus and arthropod availability across elevations and latitudes. The results demonstrated that antechinus in Queensland do not rely solely on photoperiodic cues to trigger breeding. They are able to use local environmental conditions to time breeding so that lactation coincides with timing of peak arthropod availability. Timing of antechinus breeding and flushes in arthropod availability occurred progressively later as elevation increased. More ancient Antechinus species had small ranges on mountain-tops and less flexible breeding seasons. Taken together, these results suggest that in Queensland, ancient antechinus species restricted to high elevations are likely to be most at risk from climate change, if the timing of peak arthropod availability changes. This thesis has contributed to life history, biogeography and macroecology theory at large and small spatial scales. It has enhanced our understanding of the relationship between dasyurid marsupials and their arthropod prey, and provided important information that can help us to deduce why certain dasyurids are rarer than others. ii DECLARATION BY AUTHOR This thesis is composed of my original work, and contains no material previously published or written by another person except where due reference has been made in the text. I have clearly stated the contribution by others to jointly-authored works that I have included in my thesis. I have clearly stated the contribution of others to my thesis as a whole, including statistical assistance, survey design, data analysis, significant technical procedures, professional editorial advice, financial support and any other original research work used or reported in my thesis. The content of my thesis is the result of work I have carried out since the commencement of my higher degree by research candidature and does not include a substantial part of work that has been submitted to qualify for the award of any other degree or diploma in any university or other tertiary institution. I have clearly stated which parts of my thesis, if any, have been submitted to qualify for another award. I acknowledge that an electronic copy of my thesis must be lodged with the University Library and, subject to the policy and procedures of The University of Queensland, the thesis be made available for research and study in accordance with the Copyright Act 1968 unless a period of embargo has been approved by the Dean of the Graduate School. I acknowledge that copyright of all material contained in my thesis resides with the copyright holder(s) of that material. Where appropriate I have obtained copyright permission from the copyright holder to reproduce material in this thesis and have sought permission from co-authors for any jointly authored works included in the thesis. iii LIST OF PUBLICATIONS AND PRESENTATIONS DURING CANDIDATURE Peer Reviewed Papers Collett, R.A.; Baker, A.M. and Fisher, D.O. (2018) Prey productivity and predictability drive different axes of life history variation in carnivorous marsupials. Proceedings of the Royal Society of London B: Biological Sciences, 285(1890), 20181291. Collett, R.A. and Fisher, D.O. (2017) Time-lapse camera trapping as an alternative to pitfall trapping for estimating activity of leaf litter arthropods. Ecology and Evolution, 7(18), 7527–7533. Conference Abstracts Collett, R.A. (2016) Antechinus and their prey on mountainsides. The Australian Mammal Society conference. Alice Springs, NT - September 2016. Collett, R.A. and Fisher D.O. (2017) Camera trapping as an alternative to pitfall trapping to sample prey availability of insectivorous mammals. 12th International Mammological Congress. Perth, WA - July 2017. Collett, R.A. and Fisher D.O. (2017) Rarity and resource availability for insectivorous mammals. Society of Conservation Biology conference. Cartagena, Colombia - August 2017. Collett, RA. (2018) Life history variation and extinction risk of carnivorous marsupials. UQ School of Biological Sciences postgraduate conference. Stradbroke Island, QLD - February 2018. Collett, R.A.; Baker, A.M. and Fisher, D.O. (2018) Prey productivity and predictability drive different axes of life history variation in carnivorous marsupials. The Australian Mammal Society conference. Brisbane, QLD – July 2018. iv PUBLICATIONS INCLUDED IN THIS THESIS No publications included. MANUSCRIPTS INCLUDED IN THIS THESIS Chapter 2: Collett, R.A.; Baker, A.M. and Fisher, D.O. Prey productivity and predictability drive different axes of life history variation in carnivorous marsupials. Proceedings of the Royal Society of London B: Biological Sciences. Contributor Statement of Contribution Rachael Collett Conceived the idea for the study (85%) Analysed the data (95%) Wrote the paper (80%) Diana Fisher Conceived the idea for the study (15%) Wrote and edited the paper (15%) Andrew Baker Wrote and edited the paper (5%) Simon Blomberg Advised statistical analyses and statistical paper revisions (5%) Chapter 3: Collett, R.A.; Baker, A.M. and Fisher, D.O. Winter food availability explains rarity of tropical and lowland species. Contributor Statement of Contribution Rachael Collett Conceived the idea for the study (90%) Analysed the data (100%) v Wrote the paper (85%) Diana Fisher Conceived the idea for the study (10%) Wrote and edited the paper (10%) Andrew Baker Wrote and edited the paper (5%) Eugene Mason Provided data for the study Chapter 4: Collett, R.A.; Baker, A.M. and Fisher, D.O. Do resource pulses drive latitudinal and elevational timing of breeding in the insectivorous marsupial genus Antechinus? Contributor Statement of Contribution Rachael Collett Conceived the idea for the study (90%) Analysed
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