This may be the author’s version of a work that was submitted/accepted for publication in the following source: Buckley, Thomas, Krosch, Matthew, & Leschen, Richard (2015) Evolution of New Zealand insects: summary and prospectus for future re- search. Austral Entomology, 54(1), pp. 1-27. This file was downloaded from: https://eprints.qut.edu.au/79143/ c Consult author(s) regarding copyright matters This work is covered by copyright. Unless the document is being made available under a Creative Commons Licence, you must assume that re-use is limited to personal use and that permission from the copyright owner must be obtained for all other uses. If the docu- ment is available under a Creative Commons License (or other specified license) then refer to the Licence for details of permitted re-use. It is a condition of access that users recog- nise and abide by the legal requirements associated with these rights. 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If there is any doubt, please refer to the published source. https://doi.org/10.1111/aen.12116 Austral Entomology The evo lution of New Zealand insects: summary and prospectus for future research Journal:For Austral Review Entomology Only Manuscript ID: Draft Wiley - Manuscript type: Invited Review Date Submitted by the Author: n/a Complete List of Authors: Buckley, Thomas; Landcare Research, Krosch, Mathew; The University of Queensland, Centre for Water in the Minerals Industry Leschen, Richard; Landcare Research, Keywords: Biogeography, adaptation, alpine, freshwater, island Knowledge on the evolution of the New Zealand insect fauna is reviewed and outstanding questions are highlighted. The New Zealand insect fauna is a composite of old and recent lineages and many spectacular examples of evolutionary processes are evident including; adaptive radiation, hybridisation, and unusual adaptations. We discuss the origins and evolution of four key communities within the insect fauna: terrestrial lowland insects, alpine insects, aquatic insects, and insect communities from offshore islands. Within each of these communities key lineages are discussed and in particular the key adaptations that enable these species Abstract: to thrive. Glacial history has had a dramatic impact on the New Zealand insects, and the effects on different lineages are discussed. The New Zealand insects are unique, yet many are threatened with extinction and efforts to preserve the fauna are reviewed. Despite the accumulating knowledge, major gaps still exist and these are outlined, as are opportunities to address key questions. The review ends with a synthesis and a discussion of how systematics, new technologies, and integrative approaches have the promise to improve dramatically our understanding of New Zealand insect evolution. Page 1 of 81 Austral Entomology 1 1 2 3 4 The evolution of New Zealand insects: summary and 5 6 7 prospectus for future research 8 9 10 11 12 13 14 Thomas R. Buckley 1,2,3,* 15 16 4 17 Matt Krosch 18 For Review Only 19 Richard A.B. Leschen 1 20 21 22 23 24 1 25 Landcare Research, Private Bag 92170, Auckland, New Zealand 26 27 2School of Biological Sciences, The University of Auckland, Private Bag 92019, Auckland, New 28 29 Zealand 30 31 3 32 Allan Wilson Centre, Auckland, New Zealand 33 34 4 Centre for Water in the Minerals Industry, Sustainable Minerals Institute, The University of 35 36 Queensland, Brisbane, Queensland 4072, Australia 37 38 39 40 41 *[email protected] 42 43 44 45 46 47 Running title: Evolution of New Zealand insects 48 49 50 51 52 53 54 55 56 57 58 59 60 Austral Entomology Page 2 of 81 2 1 2 3 Abstract 4 5 6 Knowledge on the evolution of the New Zealand insect fauna is reviewed and outstanding 7 8 questions are highlighted. The New Zealand insect fauna is a composite of old and recent 9 10 lineages and many spectacular examples of evolutionary processes are evident including; 11 adaptive radiation, hybridisation, and unusual adaptations. We discuss the origins and evolution 12 13 of four key communities within the insect fauna: terrestrial lowland insects, alpine insects, 14 15 aquatic insects, and insect communities from offshore islands. Within each of these communities 16 17 key lineages are discussed and in particular the key adaptations that enable these species to 18 thrive. Glacial history Forhas had a dramaticReview impact on the NewOnly Zealand insects, and the effects on 19 20 different lineages are discussed. The New Zealand insects are unique, yet many are threatened 21 22 with extinction and efforts to preserve the fauna are reviewed. Despite the accumulating 23 24 knowledge, major gaps still exist and these are outlined, as are opportunities to address key 25 26 questions. The review ends with a synthesis and a discussion of how systematics, new 27 technologies, and integrative approaches have the promise to improve dramatically our 28 29 understanding of New Zealand insect evolution. 30 31 32 33 34 Key words biogeography, adaptation, alpine, freshwater, island 35 36 37 38 39 40 INTRODUCTION 41 42 43 44 45 The origins and evolution of the New Zealand biota have been of great interest to biologists ever 46 47 since Owen (1843) began describing moa bones, and Darwin’s (1859) seminal work. Although 48 well known for the relictual lineages within the vertebrate fauna (Tennyson 2010; Allentoft & 49 50 Rawlence 2012; Mitchell et al . 2014; Rheindt et al . 2014), the New Zealand insect fauna is 51 52 equally as interesting (Fig 1). This fauna also contains relictual lineages (Leschen 2006), as well 53 54 as unusual radiations (Fleming 1975; Marshall et al . 2008), uncharacteristically depauperate 55 56 lineages (Gibbs 1980; Watt 1975), gigantism (Kuschel & Worthy 1996; Leschen & Rhode 2002; 57 Gibbs 1998; Painting et al . 2014), and many unusual ecological interactions (Dugdale 1975). In 58 59 60 Page 3 of 81 Austral Entomology 3 1 2 3 addition to understanding the evolution of particular lineages and taxa that display the above 4 5 features, the New Zealand insects offer unlimited opportunities for deepening our understanding 6 7 of fundamental evolutionary processes such as speciation (Marshall et al . 2008, 2011; Albre et 8 9 al . 2012), adaptation (Buckley & Simon 2007; McCulloch et al . 2009; Dunning et al . 2013a, 10 11 2014), hybridisation (e.g. Chinn & Gemmell 2004; Morgan-Richards & Wallis 2003; Morgan- 12 Richards et al . 2010), and phylogeography (e.g. Marske et al . 2012; Goldberg & Trewick 2011). 13 14 Previously, much research on the evolution of New Zealand insects has focussed on pattern- 15 16 based questions, such as the enduring vicariance versus dispersal debate (e.g. Craw 1988; 17 18 Trewick & Wallis 2001).For However, Review many other questio nsOnly and conundrums have been neglected, 19 which we highlight and discuss here. Biogeographic patterns in the New Zealand insect fauna 20 21 have been reviewed comprehensively at broad (e.g. Goldberg et al . 2008; Trewick et al. 2011) 22 23 and more local scales (e.g. Buckley & Leschen 2013) and we do not attempt to update these 24 25 reviews here. Our purpose is to cover outstanding questions on the evolution of the New Zealand 26 insect fauna, to summarise current knowledge, and then to outline a prospective research agenda 27 28 designed to advance the field of the evolutionary biology of New Zealand insects. Although this 29 30 review is limited to the insects, we mention other terrestrial and aquatic invertebrate taxa where 31 32 they illustrate a pattern or process that has been poorly studied in the insects, yet likely to be of 33 34 importance. 35 36 We divide this examination of the New Zealand insect fauna into several sections. The 37 38 first four sections review the origins and evolution of four different components of the insect 39 40 fauna; terrestrial insects from lowland habitats, alpine insects, freshwater insects, and insect 41 42 communities on New Zealand offshore islands. The debate on the origins of the fauna has been 43 at the heart of New Zealand insect biogeography for many years and much recent progress has 44 45 been made with phylogenetic approaches. Nevertheless, large elements of the insect biota, 46 47 particularly from the cryptic and hyperdiverse soil and saprophagous communities, have barely 48 49 been examined. 50 51 We discuss landscape changes that occurred during the Cenozoic and in particular the 52 53 reduction in land area during the Late Oligocene, postulated to have caused massive turnover in 54 55 terrestrial biodiversity (e.g. Cooper & Cooper 1995; Landis et al . 2008). Moreover, New Zealand 56 57 harbours diverse terrestrial ecosystems with numerous fascinating species interactions that are as 58 59 60 Austral Entomology Page 4 of 81 4 1 2 3 complex and intricate as on any landmass. We mention several systems and focus in particular 4 5 on the sooty mould biota, a complex web of interactions between insects, fungi and plants. The 6 7 freshwater and alpine zone insect faunas are well known for their endemism and unusual 8 9 adaptations and we discuss the origins of this diversity. We focus particularly on the unusual 10 11 physiological and biochemical adaptations that have enabled insect lineages to radiate into alpine 12 environments. These adaptations include suites of behavioural, physiological, and biochemical 13 14 strategies and we are only beginning to the scratch the surface of the underlying evolutionary 15 16 processes (Sinclair et al .