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Disentangling the drivers of ant community composition: integrating structural, spatial and inter-specific competition at multiple scales. James Schlunke A thesis submitted in fulfilment of the requirements for the degree of Doctor of Philosophy Faculty of Science University of Sydney 2016 i Abstract Understanding the drivers of community composition are crucial to providing effective conservation of biodiversity. Currently, community or ecosystem-based conservation strategies are considered the best approach for conserving cryptic or otherwise poorly quantified diversity such as invertebrate fauna, under the logic that conserving some representative subset of all communities across the landscape we are also conserving all the species that inhabit that landscape. Communities are typically defined in terms of plant community composition, and unique assemblages of plants that are rare or subject to historically high rates of loss are often afforded legal protection under biodiversity conservation legislation. This form of plant community surrogacy assumes a strong correlation between patterns of plant community composition and assemblage patterns of purportedly represented taxa, however this assumption is rarely tested. Patterns of community composition may instead be the result of a variety of other drivers besides those tied to plant community composition, including inter-specific competition or stochastic processes resulting in spatially structured assemblages. My thesis seeks to evaluate the efficacy of plant community-based strategies for conserving cryptic biodiversity, using ants as a model system. I investigate the roles of environmental and spatial drivers of ant community composition, and also the influence of inter-specific competition in shaping assemblages of ants. I first evaluated the success of a regional plant community-based conservation strategy for representation of ant assemblages across the Sydney region, Australia. I sampled ant and plant assemblage composition, and structural attributes of habitat across 40 sites from five distinct plant communities, including four listed as Endangered or Critically Endangered under state and national legislation. I found a weak association of ant assemblages with plant community composition, with only those community pairs varying strongly in structural composition of vegetation supporting unique i assemblages. Despite the weak associations, plant community composition was the strongest predictor of ant assemblages, however structural variation amongst samples also contributed to observed assemblage composition. Spatial turnover amongst ants was low overall, however I found stronger within-community turnover patterns, and the strength of this relationship also varied greatly amongst community types. I then investigated the relative importance of plant community, habitat structure and spatial drivers of ant assemblage composition at fine scales in a mosaic of plant communities within the NSW South-West Slopes bioregion. I used Multiple Regression on Distance matrices to partition explained variation amongst assemblage drivers. Spatial turnover was the strongest predictor of ant assemblage composition, with assemblages also showing some association with habitat structure. Plant community composition alone showed weak significant association with ant assemblages, however this association was driven entirely by shared spatial autocorrelation of assemblages and structural differences amongst plant communities. Habitat complexity was found to influence assemblages by reducing the fine-scale abundances of highly aggressive competitively dominant species. Abundances of dominant species at fine-scales in turn negatively affected the abundances of competitively subordinate ant species. The abundance of dominant species had a negative effect on species richness of subordinate species at fine scales, however they had no effect at intermediate or course spatial scales. I found competitive exclusion between the two most abundant dominant species in the study area, Iridomyrmex purpureus and I. chasei. To investigate the potential influence of variable competitive influences on ant community assemblages, I sampled ant assemblages along well-defined territory boundaries. I also compared the strength and uniformity of competitive dominance at food resources within territories of the two species. I found very strong inter- specific competition between dominant species, resulting in rapidly shifting territory ii boundaries with I chasei consistently encroaching into the territory of I. purpureus and regularly usurping nests. Food resources were more consistently won by the numerically dominant I. chasei, resulting in lower species richness of affiliate ant species attending baits, and significantly different assemblage composition of species attending baits between territories occupied by the two species. In conclusion, I found that plant community surrogacy provided only weak representation of ant assemblages, and only at course scales when comparing broadly different plant community classes. Within communities, and amongst structurally similar plant communities, spatial turnover in assemblages was strong and resulted in highly spatially structured ant communities. Assemblages also appear to be structured in part by inter- specific competition, both in terms of non-random patterns of spatial association amongst dominant species, but also through suppression of competitively subordinate species by dominant ants. These finding are especially important to the implementation of biodiversity offsetting strategies, as ant assemblages represented in compensatory habitat would likely be compositionally distinct from those lost through development if offsets are located large distances from lost habitat. iii Declaration The research described in this thesis, except where acknowledged, is all my own original work. Conceptual ideas were developed in collaboration with my supervisors Associate Professor Dieter Hochuli and Dr Mathew Crowther. Signed: Name: James Schlunke iv Acknowledgements I would first like to thank my two supervisors, Dieter Hochuli and Mathew Crowther. I’ve had the pleasure of knowing you both for ten years, and not only have you been a truly exceptional and unique supervisory team, you’ve both also shaped my whole outlook as an ecologist. I’ll especially miss the seemingly endless hours having coffee and our occasional BLOBS meetings. Dieter, without your 2nd year undergraduate Entomology course I almost certainly wouldn’t have taken the route I have, and I’ve never once regretted it. Truly no other lab has the same dynamic and that can perhaps be attributed to your finely tuned sense of humour and welcoming nature, and to a lesser degree to your sharp dress sense, eclectic musical tastes and encyclopaedic knowledge of pop culture. Thanks for welcoming me into the lab for Honours back in 2007 and letting me stick around as long as you have. May your flowing, luxuriant hair forever remain as full-bodied as it is today. Mat, thanks for being an endless source of help, advice, inspiration, distraction, amusement and perhaps procrastination. Your technical and help has been absolutely invaluable and many of the skills I’ve developed over the last five years I owe to you. Thanks for always pushing the boundaries of both science and public acceptability. I’d like to extend an absolutely enormous thankyou to all the Hochuli lab members past and present for always making my time an enjoyable and enriching experience. You’re a unique bunch of people and it’s been laughs the whole time. Fran van den Berg, truly nobody else could empathise as much as you in this whole thing. Your endless hours, weeks, months and even years tending spiders to adulthood were a constant source of inspiration to work harder. You’ve made this fun throughout the best and worst of times, and you’ve both enabled me to stick around far beyond my use-by-date, and also been an inspiration to finish once you finally did. Thanks to all the current postgrads Petah Low, Lizzy Lowe, Jo Haddock, Ryan Leonard, Jayne Handford, Henry Lydecker, Manuel Lequerica, Lucy Taylor, postdocs v Caragh Threlfall and Erin Leckey, and also to all the Honours students who’ve come through the lab in all this time. Thanks also to all the friends in the postgrad community and in room 408, especially Tony Popic for the laughs and enabling my procrastination early on, and Lindsey Gray for the fun times and sneaky lunchtime beers. Thanks to Alan Andersen for your invaluable help with ant identifications, very helpful advice and generous hospitality. Thanks also to all the staff and students at the CSIRO Tropical Ecosystems Research Centre for being so welcoming, accommodating and helpful during my stay. Thankyou to David Schlunke and Jeanette Gash for being so helpful and accommodating when I was back home in the field. Thanks especially for all the Chinese meals in Temora when I was around. Ultimately I wouldn’t have the passion for nature I have if not for my dad, and having the chance to study ecology in the same place I grew up has been fantastic. I doubt many PhD students get to walk to their study sites from their family home. And thanks also to the rest of my family Anna, Eva and Lynette (mum) for your help and support throughout. Thanks also to the people who helped me in the field, both for this project and my prior ill- fated foray into the world of sexually-deceptive orchids; George Madani, Trevor Wilson, Matt Renner, James Bevan,
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